D.M.R.G. Mayakaduwa1 and T.D. Silva1 1Department of Plant Sciences, University of Colombo, Colombo 03, Sri Lanka Corresponding authors e-mail [email protected] Tel 94775086752 Flow cytometric detection of haploids, diploids and mixoploids among the anther-derived plants in Indica rice (Oryza sativa L.) reproduced in the clustering patterns obtained from morphometric analysis Abstract Keywords Anther culture-callus induction-green plant regeneration-stomatal measurements Introduction Anther culture technique can be used to produce homozygous lines immediately, thereby shortening considerably the time required for inbred line development in plants by conventional methods. For the technique to be effective, plants that are regenerated from anther culture should be haploids or dihaploids (doubled haploids). Since regenerated plants do not always have the expected ploidy, an assessment of the level of ploidy of anther-derived plants is often required. A direct method of establishing ploidy is to obtain chromosome counts from regenerated plant tissue. However, preparation of metaphase chromosome spreads for chromosome counting from mitotic cell nuclei is ineffective due to the small size and poor staining of the chromosomes. Therefore, indirect methods such as cell size measurements that can be applied more easily have been employed to estimate ploidy in plant tissue, based on the general premise that there is an overall increase in cell size with increase in ploidy. Stomatal size (length /width), stomatal density (Borrino and Powell 1988 Silva et al. 2000 Mohommadi et al. 2012 Shrestha and Kang 2016), as well as chloroplast number in guard cells (Ho et al. 1990) have been shown to correlate well with chromosome number or ploidy level of many different plant species. On the other hand flow cytometry, which determines the DNA content in individual cell nuclei with reference to a known standard, is a rapid and accurate method that is easily applied at individual cell level. This method is most useful in analyzing plants derived from in vitro culture because it allows detection of chimeras which is not an uncommon phenomenon in cell cultures. Therefore, indirect estimates of chromosome number in regenerated plants should be coupled with flow cytometry to obtain a comprehensive view of the ploidy in plants that are regenerated from tissue culture. In the present study, the ploidy status of anther-derived plants of Indica rice variety At 303 was examined using flow cytometry. The possibility of using indirect methods such as stomatal and other morphological trait measurements to discriminate different levels of ploidy was also examined. Methodology Anther culture in Indica rice var. At 303 Plant material Mother plants of the Indica rice variety At 303were raised in a field at Regional Rice Research and Development Centre, Bombuwela, Sri Lanka, under standard agronomic practices. When plants reached the booting stage they were uprooted and transplanted in pots with paddy soil and brought to the laboratory of Department of Plant Sciences, University of Colombo. The pots were kept outside the laboratory until panicles were ready for harvest. Immature panicles were harvested while panicles were still concealed within leaf sheath, to obtain anthers with microspores at the appropriate stage of maturity (late uni-nucleate) for culture (Mayakaduwa and Silva 2017). Panicles were collected from mother plants between 9.00 10.00 am. Anther culture and callus induction Harvested panicles were cold pre-treated at 10 0C for 7 10 days. Cold pre-treated panicles were surface disinfected according standard procedures (Silva and Ratnayake 2009). Anthers excised from spikelets were cultured on N6 callus induction medium (Chu 1978), modified by replacing sucrose with maltose (60.0 g L-1). Callusing medium was supplemented with the growth regulators, 2,4-Dichlorophenoxyacetic acid (2,4-D) 0.5 mg L-1, Naphthaleneacetic acid (NAA) 2.5 mg L-1, and Kinetin (0.5 mg L-1) and solidified with agar (8.0 mg L-1). Up to 200 anthers were placed on a single Petri dish containing 15 mL of the culture medium. Culture plates were incubated in the dark at 272 0C for callus induction. Plant regeneration and acclimatization Analysis of anther-derived rice plants Flow cytometric analysis Leaf samples of ten fully grown plants regenerated from anther culture were analyzed by flow cytometry to establish their ploidy. Leaves of normal diploid plants of similar maturity, obtained from the same rice variety, were also analyzed and used as the standard for 2C DNA content. Samples were processed using Cystain UV Precise P Kit (Partec GmbH, Germany) and DAPI stained nuclei were analyzed on flow cytometer CyFlow Space (Partec GmbH, Germany). Stomatal measurements Stomatal measurements were obtained from six plants of rice variety At 303 regenerated from anther culture and grown to maturity. Epidermal peels were obtained from the two youngest leaves to study stomata. From each leaf 5 lower epidermal peels were observed under the light microscope and in each peel 5 stomatal units were selected randomly to obtain length (distance between the ends of the two subsidiary cells measured lengthwise) and width (horizontal distance across the two subsidiary cells at the widest point). Measurements were made under X 100 magnification using an oil immersion lens. To determine the stomatal density in each epidermal peel, the number of stomata observable in a microscopic field under X 10 magnification was counted first and converted to the number of stomata per unit area. Micrographs were taken using a light microscope (Carl Zeiss, Oberkochen, Germany) with a Axiocam digital camera (Carl Zeiss) and images were processed with Zen 2012 (blue edition) imaging soft-ware. The above measurements were repeated on the six diploid plants of the same rice variety (At 303) raised in vitro from seed culture and grown to the same level of maturity for comparison. Morphological and reproductive traits Measurements were obtained for plant height, leaf blade length and width, number of reproductive tillers, number of spikelets per panicle and panicle length. The data were recorded for the six anther-derived and six seed grown plants separately. Data analysis Data on stomatal length, width and density were analyzed using one-way ANOVA with SAS Release 9.0. Mean comparisons were carried out using Tukeys multiple comparison test. Hierarchical cluster analysis was performed with stomatal data as well as morphological and reproductive trait measurements using SPSS 16.0. Two separate dendograms, one for stomatal measurements and the other for morphological data, were developed based on the similarity relationships. Results and Discussionand Methodsy external marker as the guid eto sativa L.) PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 PAGE ARABIC 0 Anther culture in Indica rice variety At 303 Callus induction occurred from cultured anthers approximately 4 weeks after anther inoculation. Of 631 anthers cultured, 24 anthers responded (4.12) by producing calli which is a reasonable rate of success for the rice variety At 303 (Mayakaduwa and Silva 2017). After transfer to the regeneration medium 8 calluses (33) produced green shoots and all of these were moved to pots for acclimatization (Fig. 1a-b). The plants that survived the acclimatization were grown to maturity (Fig. 1 c). The mature plants were characterized by flow cytometry and analysis of stomatal data together with morphological and reproductive traits. Fig. 1 Acclimatization of anther-derived plants of rice variety At 303 a- green shoots with roots regenerated from anther culture, b- plantlets in pots containing sand being acclimatized inside propagators, c- plantlets moved to pots with paddy soil and grown to maturity in a glass house (bar a 3.0 cm, bar b 5.0 cm, bar c 15.0 cm) Analysis of anther-derived rice plants Flow cytometry Flow cytometric analysis of regenerated plants showed that they could be haploids, diploids or mixoploids, as illustrated by the representative histograms (Fig. 2). Of the 10 plants, 4 were determined to be haploid (1C), 5 diploid (2C) and 1 plant was a mixoploid (1C and 2C). Presence of haploid plants among the regenerants confirmed their microspore origin. In the diploids and mixoploid, cells with higher ploidy may have arisen from doubling of chromosomes in vitro. However, further confirmation will be required using molecular markers to establish that their origin was from haploid microspores and not from diploid somatic tissue of anther wall. Fig. 2 Flow cytometric analysis of nuclear DNA content in anther-derived plants against seed-derived normal diploid plants (variety At 303) stained with DAPI a- haploid plant with G1 DNA peak (1C) set at channel number 100, b-diploid plant with G1 DNA peak (2C) set at channel number 200, c-mixoploid plant with two G1 DNA peaks (1C and 2C) at channel numbers 100 and 200 Stomatal measurements Stomatal size and density have been used as indirect measurements to determine ploidy of plants. In general plants with higher ploidy have larger cell sizes and lower stomatal densities compared to plants with lower ploidy status (Dhooghe et al. 2011 Shrestha and Kang 2016). Table 1 Stomatal measurements of anther-derived plants and plants grown from seed (variety At 303) Clustering based on the stomatal measurements produced two main groups. Seed-derived normal diploid plants and plant nos. 7, 8, and 10 of anther-derived plants formed a single distinct cluster (at a relative distance 0.97). Plant no. 9 (anther-derived) also showed a closer relationship to this group although at a relative distance 2.33. Plant nos. 11 and 12 separated out with the greatest distance from other phenotypes (at a relative distance 25) indicating their genetic uniqueness (Fig. 4). Therefore, it may be assumed that these two plants which had the smallest cell size and highest stomatal density are true haploids whereas the remaining anther-derived plants which clustered together with normal diploid plants are diploid or may even be mixoploid. Fig. 4 Dendrogram showing clustered relationships of the seed-derived and anther- derived plants of the variety At 303, analysed based on stomatal measurements sseed-derived plants aanther-derived plants Morphological and reproductive traits The seed plants, on average scored higher values in all the measured morphological parameters except in plant height (Table 2). The two anther-derived plants 9 and 10 were the taller (79.8 cm and 69.8 cm respectively) than the normal seed plants (62.1 cm 68.5 cm) while two other anther-derived plants 11 and 12 were distinctly shorter (45.2 cm and 46.8 cm respectively) than all others. The latter two plants scored lower values for all other parameters including leaf blade length and width, panicle length and number of spikelets per panicle. The clustering pattern in the dendogram developed using morphological data was similar to the one observed with stomatal data. However, morphological data produced three clusters. All seed plants were contained in one cluster. Anther-derived plants separated into two clusters, one that included plants 7, 8, 9 10, and another distinctly separate cluster with plants 11 and 12 at relative distance 25 (Fig. 5). The greatest distance displayed by plants 11 and 12 coupled with significant differences observed with Tukeys mean comparison tests on stomatal measurements further strengthen the suggestion that these two plants have a haploid genome. The other anther-derived plants which showed a closer affinity to seed grown normal diploids maybe diploids or mixoploids. Table 2 Morphological and reproductive trait measurements of anther-derived plants and seed-derived plants at the panicle emergence maturity (variety At 303) Plant NumberPlant typePlant height (cm)Leaf blade length (cm)Leaf blade width (cm)Number of reproductive tillers/plantPanicle length (cm)Number of spikelets/ panicle1seed-derived68.534.21.7214.5622seed-derived65.433.21.5214.0703seed-derived63.028.61.5115.4584seed-derived62.129.81.5115.1615seed-derived62.334.01.4214.8546seed-derived64.537.81.5215.6507anther-derived66. Fig. 5 Dendrogram showing clustered relationships the seed-derived and anther- derived plants of the variety At 303, analysed based on morphological trait measurements sseed-derived plants aanther-derived plants Flow cytometric analysis revealed that plants regenerated from anther culture of Indica rice variety At 303 could be haploid, diploid or mixoploid. Stomatal measurements and other morphological data also corroborated the flow cytometric evidence. In the plants subjected to flow cytometry, only 4 of 10 plants (40) were haploid. Analysis of six regenerated plants using stomata and other morphological parameters indicated that a similar proportion (33) were likely to be haploid. Thus it would appear that of the total regenerants less than half are haploids. Anther culture is performed with the intention of regenerating haploid or doubled haploid plants. Therefore in a system where majority of the regenerants are non-haploids, it is necessary to understand the cellular origin of these plants. Flow cytometric analysis of the regenerated plants determined that 50 of plants were diploid. It could be that these were doubled haploids. Endoreduplication of chromosomes leading to spontaneous chromosome doubling is a relatively common phenomenon in in vitro culture systems (Ochatt et al. 2011) and some systems rely on it to produce doubled haploids or homozygous lines. However the origin of doubled haploids needs to be substantiated by analyzing genetic loci for homozygosity using molecular methods since diploids can also arise from regeneration of maternal anther tissue. In several studies, SSR markers have been used to characterize plants produced through anther culture in rice (Sah and Niroula 2007 Luitel et al. 2012 Rout et al. 2016). However, molecular characterization was not attempted in the present analysis due to various constraints. Thus, further determination of the origin of the diploid plants cannot be concluded. A small proportion (10) of plants analyzed by flow cytometry was mixoploid. It was confirmed that mixoploids contained haploid and diploid cells but no higher ploidy levels. Presence of haploid cell lines in mixoploids can be considered as lending support to the hypothesis that plants originated from microspores but during the initial phase of cell division there was endoreduplication of chromosomes leading to some cell lines becoming diploid. Presence of mixoploids in plants regenerated through anther culture has been reported earlier in rice (Cha-um et al. 2009). Nevertheless, mixoploids are undesirable in a system of dihaploid production because they would be partially sterile and genetically unstable. In recent studies on anther culture of Indica rice, ploidy of regenerated plants have been determined on the basis of visual observations on morphology and fertility (Sah and Niroula 2007 Mishra et al. 2015). Haploid plants were identified by being small and sterile compared to diploids that showed normal growth and fertility. Similar observations were made in the present study with regard to morphological data. In addition to such morphological traits, indirect methods to establish ploidy, especially the use of stomatal measurements (length, width and density) were examined. These cell measurements have been accepted as being useful in estimating the ploidy in many different plant species (Kadota and Niimi 2004 Sun et al. 2009 Yan et al. 2016). Although these cell measurements were sufficiently robust in discriminating haploids from diploids, the use of stomatal measurements may not allow adequate separation between diploids and mixoploids. Thus it could be suggested that while stomatal size and density measurements could be used to provide a reasonable guide to ploidy determination of plants regenerated from indica rice anther culture, the precise determination of ploidy will have to rely on chromosome counting or flow cytometry. Conclusions Determination of ploidy of regenerated plants derived from anther culture can be considered as essential as the anther culture technique is performed specifically for the production of homozygous lines. This study reveals that different levels of ploidy existed among the regenerated plants from anther culture, as detected by flow cytometry. Further, the ploidy estimates from stomatal, morphological and reproductive traits analyses highly corroborated the flow cytometric evidence. Therefore this study validates the use of flow cytometry combined with stomatal measurements, morphological and reproductive traits for successful characterization of ploidy status of anther-derived plants in Indica rice. Further, this suggests the importance of improving the efficiency of regenerating haploids or dihaploids through anther culture in Indica, in order to apply the anther culture technique as a practically viable breeding tool. Acknowledgement Funding by the National Science Foundation (NSF) of Sri Lanka under the research grant number RG/2011/BT/10 is gratefully acknowledged. Assistance provided by the Regional Rice Research and Development Centre, Bombuwela, Kalutara, Sri Lanka is also appreciated. Literature cited Borrinoe, M. and W. Powell (1988). Stomatal guard cell length as an indicator of ploidy in microspore-derived plants of barley. Genome 30 158 160. Cha-um, S., B. Srianan, A. Pichakum, and C. Kirdmanee (2009). An efficient procedure for embryogenic callus induction and double haploid plant regeneration through anther culture of Thai aromatic rice (Oryza sativa L. subsp. indica). In Vitro Cellular and Developmental Biology Plant. 45 171179. Chu, C.C. (1978). The N6 medium and its applications to anther culture of cereal crops. Proceedings of the Symposium on Plant Tissue Culture. Science Press, Beijing, pp. 4350. Dhooghe, E., K. Van Laere, T. Eeckhaut, L. Leus, and J. Van Huylenbroeck (2011). Mitotic chromosome doubling of plant tissues in vitro. Plant Cell Tissue and Organ Culture. 104 359373. Ho, I., Y. Wan, J.M. Widholm, and A.L. Rayburn (1990). The Use of stomatal chloroplast number for rapid determination of ploidy level in Maize. Plant Breeding 105 203210. Kadota, M., and Y. Niimi (2004). Production of triploid plants of Japanese pear (Pyrus pyrifolia Nakai) by anther culture. Euphytica. 138 141147. Luitel, B.P., P.B. Adhikari, S.L. Shrestha, and W. Kang (2012). Morphological characterization of anther derived plants in Minipaprika (Capsicum annuum L.). Korean Journal of Breeding Science. 44 (4) 450461. Mayakaduwa, D.M.R.G., and T.D. Silva (2017). A cytological indicator allows rapid assessment of microspore maturity, leading to improved in vitro anther response in Indica rice (Oryza sativa L.). In Vitro Cellular and Development Biology Plant. 53 (6) 591597. Mishra, R., G.J. Narashima, R. Nageswara, and P. Kaushal (2015). 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Kang (2016). Stomata length and density as an indicator of ploidy level in Sweet Pepper (Capsicum annuum L.). Research Journal of Recent Sciences. 5 410. Silva, P.A.K.X.M., S. Jacques, and M.H. Zanettini (2000). Induction and identification of polyploids in Cattleya intermedia Lindl. (Orchidaceae) by in vitro techniques. Ciencia Rural, Santa Maria. 30 (1) 105111. Silva T.D., and W.J. Ratnayake (2009). Anther culture potential of indica rice varieties, Kurulu thuda and Bg 250. Tropical Agricultural Research and Extension. 12 (2) 5356. Sun Q., H. Sun, L. Li, and R.L. Bell (2009). In vitro colchicine-induced polyploid plantlet production and regeneration from leaf explants of the diploid pear (Pyrus communis L.) cultivar, Fertility. Journal of Horticultural Science Biotechnology. 84 (5) 548552. Yan, H.J., Y. Xiong, H.Y. Zhang, and M.L. He (2016). In vitro induction and morphological characteristics of octoploid plants in Pogostemon cablin. Breeding Science. 66 169174. 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