Congenial in Vitro γ-Ray Induced Mutagenesis Underlying the Varied Array of Petal Colours in Chrysanthemum (Dendranthemum Grandiflorum Kitam). ‘Candid’

doi:10.21203/rs.3.rs-617238/v1

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Congenial in Vitro γ-Ray Induced Mutagenesis Underlying the Varied Array of Petal Colours in Chrysanthemum (Dendranthemum Grandiflorum Kitam). ‘Candid’

https://doi.org/10.21203/rs.3.rs-617238/v1

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Chrysanthemum (Dendranthemum grandiflorum kitam.) is amongst the top ten cut flowers globally. The flower colour of ancestral species is restricted to white, yellow, and pink and is acquired from anthocyanins, carotenoids, and the dearth of both pigments, respectively. An extensive array of flower colours, like orange, dark red, purplish-red, and red, has been bred by enhancing the variety of pigments or the amalgamation of both pigments. In recent times, green-flowered cultivars having chlorophylls in their ray petals have been produced and have grown a reputation. Furthermore, violet /blue flowers have been bred via transgenic interventions. Flower colour is considered as critically acclaimed feature of any flower cultivar especially chrysanthemum. Creating newer chrysanthemum cultivars with novel features, for instance, new flower colours in a time and input optimised approach, is the eventual ambition for breeders. Exploring the molecular mechanisms that control flower pigmentation may present imperative suggestions for the rational manoeuvring of flower colour. To generate a diverse array of flower colour mutants in chrysanthemum cv. “Candid” through mutagenesis, in vitro grown micro shoots were exposed to 10, 20, 30, and 40 Gy gamma irradiation at 100 Gy per minute and were evaluated for different parameters. The rhizogenesis parameters declined with the increase in irradiation dose from 0 Gy to 40 Gy, while as, 10 Gy dose proved to record minimum decline in contrast to the control. Survival, leaf size, and the number of leaves plant^− 1 after the 8th -week interval also decreased with the increasing trend of gamma irradiation dose but recorded a minimum decline in plants raised from shoots irradiated with 10 Gy gamma irradiation dose with respect to the control. Apparently, the minimum delay in the number of days to floral bud appearance took under 10 Gy compared to control. The highest number of flower colour mutants were recorded under 10 Gy (light pink, orange-pink, white and yellow). Demountable mutation frequency based on flower colour was desirable in plants irradiated with the slightest dose of 10 Gy

Plant Physiology and Morphology

Plant Molecular Biology and Genetics

Chrysanthemum

colour mutants

mutagenesis

gamma irradiation

mutants

Chrysanthemum is a prevalent and essential cut flower crop grown worldwide in China, Japan, France, the USA, India, and the UK. It is a keyfloricultural crop, and ranks 2nd in terms of cut flowers following rose, amongst the floricultural crops traded in the international flower market (Kumar et al. 2006). The complex genetic hetrozygosity makes the cultivated chrysanthemum an unlimited source of new flower form and cultivars. The common garden chrysanthemum is hexaploid with 54 chromosomes (Wolff 1996). It is propagated vegetatively and has a robust self-incompatibility system (Richards 1986); therefore, new varieties are complicated to acquire by crossing. Usually, new varieties have been developed from spontaneous mutations in vegetative reproduction, sports, being some variations more stable than others (Miñano et al., 2009). In recent years induced mutations along with the somaclonal variations resulting from the tissue culture practices have been in work as an innovative source of variability (Schum 2003, Datta et al. 2005, Jain et al. 2006, Zalewskam and Miller 2007, Chatterjee et al. 2006, Barakat et al. 2010) Even though comprehensive efforts have been undertaken to expand uniqueness in chrysanthemum cultivars by induced mutations employing physical and chemical mutagens (Broertjes and Van-Harten 1978), there is a constant need to investigate the chances of a new cultivars for horticulture trade. Mutation breeding by radiation has been extensively used to improve well-established plant cultivars and develop novel variants within superior consumer characteristics. While the majority of cultivated chrysanthemum varieties are polyploids with huge genetic heterogeneity, mutants with allied flower shape, floral size, and colour, are frequently recovered. Associated flower colours with chimeric tissue can be simply stimulated by radiation and isolated with in vitro apparatus (Kumar et al., 2006). Mutation procedures are used because chrysanthemum is a hexaploid plant and vegetative propagated, making it difficult to conduct the hybridization (Broertjes and Van-Harten 1978). Genetic variation is essential in any plant breeding programme for crop improvement. Mutation breeding is an efficient way to produce heritable change, particularly for the flower colour. Increasing demand for the new form of chrysanthemum leads to research for obtaining new varieties. Mutation breeding by radiation, a farming appliance of nuclear technology, and has been widely exploited to develop well-adapted plant varieties by one or a few important qualities (Kumar et al. 2006, Jain and Spencer (2006, Dwimahyani and Widiarsih 2010, Chatterjee et al. 2006) Commercially important traits in horticulture plants have been altered positively by the various physical mutagens. Gamma rays are widely used among the physical mutagens for inducing mutations in flowering plants due to their easy application and high efficiency. The physical irradiations have been used efficiently to induce mutation in chrysanthemum and the optimal dose range from 1.0 to 3.0 Krads correlated to the type of genotype (Dilta et al., 2003). While going for mutation breeding programmed various factors like choice of material, character to be improved, type of mutagens and its dose to be used, an experimental procedure to be chosen should be considered. Thus through mutation breeding, it is possible to induce a genetic variation for quantitative and qualitative characters that is heritable of sufficient magnitude and frequency of interest in the breeding programme. Accordingly, the genetic variability produced by mutation was studied to develop a novel cultivar in chrysanthemum having momentous customer fondness. Consequently, with deliberation to above facts, the current study “Congenial in vitro γ- ray induced Mutagenesis underlying the diverse array of petal colours in chrysanthemum (Dendranthemum grandiflorum kitam) cv. ‘Candid’ ” was undertaken with an objective to generate a diverse array of flower colour mutants through mutagenesis.

Tissue culture developed micro shootlets of chrysanthemum cv. ‘Candid’ (parent cultivar, Figure-1) were exposed to Cobalt⁶⁰ gamma irradiation doses of 0, 10, 20, 30, and 40 Gy at 100 Gy per minute and were allowed to raise vegetatively mutated generations first and second at 5-week intervals. Finally, shoots obtained from vegetatively mutated generation 2 were allowed for rooting and consequent acclimatization. Rooted shoots were allowed to grow in pots in the field to obtain new enviable colour mutants, and rooting parameters were recorded in terms of percentage rooting and the number of roots per shoot. Survival (%), leaf area plant^− 1 (cm²), number of leaves plant^− 1 were recorded at 4th and 8th weeks growth in the field. Days to flower bud appearance were recorded at the initiation of flower bud appearance. Plant height was noted at the end of full flower bloom. Flower colour was recorded in terms of the difference between the parent flower and mutants obtained. The frequency of mutation was calculated based on flower colour, as the ratio between such desired or undesired colour mutant and whole plants irradiated with each gamma irradiation dose.

The investigation was carried out in completely randomised design with four replications and the data composed on various observations during the trail was preceded to analysis of variance (ANOVA). To assure model assumptions for ANOVA percent age data were subjected to square root transformation as recommended by Steel and Torrie (1980) and the average data was classified by Duncan’s multiple range test (Gomez & Gomez, 1983).

The irradiation doses had an enervating effect on all the rooting parameters in comparison to control (Table 1). A significant decrease in mean rooting percent and the number of roots shoot^− 1 in all the irradiation treatments compared to control was observed. Among irradiation treatments, the minimum decline in rooting and number of roots were recorded under 10 Gy dose, followed by 20 and 30 Gy irradiation doses. Maximum decline in rooting and the number of roots were registered with 40 Gy dose (Fig. 2-a, and b, Fig. 1, 3a and 3b). Rhizogenesis is a procedure of dedifferentiation of definite pre-determined cells near the vascular bundles. Any harm to cell division capability will have an unconstructive outcome on the dedifferentiation of cells and consequent reorganization into root primordia. This could cause malfunction of rooting or deferred emergence of roots. Singh et al. 1999 also reported that increased doses of gamma irradiation (from 20 to 50 Gy) decreased the rooting percentage of carnation cv. ‘Espana.’ Radiation treatments also delayed root initiation significantly in comparison to control. Sooch et al.2000 observed delayed root initiation of carnation shoots of cv. ‘Scania’ under 1.00, 1.50, and 2.00 K-rads gamma irradiation doses. The deleterious effects of radiations also showed a significant decline in root number per shoot under 10 to 30 Gy treatments. El-sharnouby and El-Khateeb 2005 also reported that most of the gamma irradiation treatments (10, 20, 30, and 40 Gy) without or with NAA in the rooting medium decreased the number and the length of roots in the carnation cultivars “Medea,” “Candela” and “Picaro.” All the above-quoted studies seem closer to the findings recorded in the present study. Survival of rooted shoots at the end of 4 weeks was significantly minimum by the shoots treated with 40 Gy dose as against control, followed by 30 and 20 Gy dose (Table 2). Under a minimum dose of 10 Gy, there was a minimum decline in survival of shoots at the end of the 4 weeks over control. At the end of 8 weeks, shoots treated with 10 Gy dose recorded maximum survival, followed by 20 and 30 Gy dose (Fig. 2, 3c and 3d). While lowest survival per cent was recorded in 40 Gy irradiated shootlets corresponding to a sharp decline compared to control.

Table 1

Influence of ⁶⁰Co gamma irradiation on rhizogenesis in shoots of Chrysanthemum (*Dendranthemum morifolium* L.) cv. “Candid”
Dose	Rooting (%)	Root number shoot^− 1
0 Gy	89.28a ± 2.38 (9.44) *	15.75a ± 0.50 -
10 Gy	85.71 b ± 0.00 (9.26) * (1.90) **	7.75 b ± 0.96 (50.79) **
20 Gy	79.76c ± 2.38 (8.93) (5.40)	5.50 c ± 0.58 (65.07)
30 Gy	74.99 d ± 2.38 (8.66) (8.26)	3.00d ± 0.00 (80.95)
40 Gy	73.80d ± 2.75 (8.59) (9.00)	2.75d ± 0.50 (82.53)
L.S.D _P≤0.05	0.18	0.90
Figures in the parenthesis are the square root transformed values of percentage data *Figures in the parenthesis are the per cent decrease in vegetative parameters over control

Table 2

Influence of ⁶⁰Co gamma irradiation on survival of rooted plantlets under polyhouse conditions in Chrysanthemum (*Dendranthemum morifolium* L.) cv. “Candid”
Dose	Survival (%)
Dose	Week 4th	Week 8th
0 Gy	95.23a ± 0.00 (9.76)*	95.23a ± 0.00 (9.76)*
10 Gy	85.71b ± 3.89 (9.25)* (5.22)**	82.14 b ± 2.38 (9.06)* (7.17)**
20 Gy	76.18c ±3.89 (8.72) (10.65)	67.85c ± 4.56 (8.23) (15.67)
30 Gy	61.90d ± 3.89 (7.86) (19.46)	53.56d ± 4.56 (7.31) (25.10)
40 Gy	53.56e ± 5.99 (7.31) (25.10)	40.47e ± 6.15 (6.34) (35.04)
LSD _(P≤0.05)	0.17	0.27
Figures in the parenthesis are the square root transformed values of percentage data *Figures in the parenthesis are the per cent decrease in vegetative parameters over control

Broertjes and Lock (1984) obtained 100% survival when chrysanthemum plantlets transferred to soil were irradiated with 2.5 or 5 kGy. The deleterious chimera load carried by the plants leads to mortality in post-irradiation proliferative generations. Another reason might be the formation of the low or reduced wax component on the post-irradiation plants. The wax module decides the pace of water loss through the cuticle and the vulnerability of tissue-cultured plants to desiccation accredited to a decline or lack of wax acting as an antitranspirant. The epicuticular wax is reduced or absent on the carnation leaves of in vitro cultured plants compare to glasshouse or field-grown plants (Sutter and Langhans1979). Still, during acclimatization, the density of waxes boosts as the humidity recedes (Wardle et al. 1983). Since the irradiation impairs the plants' epidermal skin, low wax formation during the acclimatization process leads to mortality.

Influence of γ- rays on leaf area and number of leaves

Gamma irradiated treatments significantly recorded a decline in leaf number plant^− 1 and leaf size in both the intervals, i.e., 4 and 8 weeks, compared to control (Table 3). At the end of 4 weeks, significantly minimum leaf number plant^− 1 and size were registered under the highest dose of 40 Gy, followed by 30 and 20 Gy. The lowest gamma irradiation dose, 10 Gy, recorded a minimum decrease in leaf number and size compared to the control. At the end of 8 weeks, both leaf number as well as leaf size improved in all the gamma irradiation doses, including the control plants but recorded a similar trend of decline in both the parameters as in the 4-week interval with the successive gamma irradiation doses (Fig. 3e to 3h).

Table 3

Influence of ⁶⁰Co gamma irradiation on leaf number and leaf size in Chrysanthemum (*Dendranthemum morifolium* L.) cv. “Candid”
Dose	Leaf number plant^− 1		Leaf size (length/width) (cm²)
Dose	4th week	8th week	4th week	8th week
0 Gy	14.00a ± 0.82	15.75a ± 0.96	22.31a ± 3.30	28.52a ± 1.18
10 Gy	11.00b ± 0.82 (21.42)	13.00 b ± 0.82 (17.46)	20.42 a ± 1.45 (8.43)	27.80 a ± 0.89 (2.52)
20 Gy	9.25 c ± 0.50 (33.92)	11.75 bc ± 0.96 (25.39)	17.34 b ± 01.32 (22.24)	20.42 b ± 1.45 (28.40)
30 Gy	8.75 cd ± 0.50 (37.50)	10.75 cd ± 0.96 (31.74)	13.52 c ± 0.72 (39.37)	14.82 c ± 1.23 (48.03)
40 Gy	8.00 d ± 0.00 (42.85)	9.75 d ± 0.96 (38.09)	5.31 d ± 0.66 (76.18)	10.74 d ± 2.14 (62.34)
L.S.D _P≤0.05	0.92	1.41	2.69	2.37
Figures in the parenthesis are the per cent decrease in vegetative parameters over control

Leaf area increment results from the growth of cells mainly controlled by growth regulators (auxins). Higher exposure to gamma irradiation agitate synthesis of auxins, hence leads to decreased leaf area. Simard et al.1992 and Cassels et al. 1993 recorded biological damage in carnation on increasing the dose of radiation. Misra and Bajpai, 1983a in gladiolus; Gupta et al., 1982 in costus; Gupta et al., 1974 in tuberose; Acharya and Tiwari 1996; Siranut et al., 2000 in chrysanthemum; Srivastava et al. 2007 in gladiolus; Misra et al. 2009 in chrysanthemum and Kahrizi et al. 2011 in rose also accounted the decrease in number of leaves with the raise in dosage of gamma irradiation whereas, Kumari et al. 2013 reported decline in leaf size in terms of length and width of plants treated with higher doses of gamma rays in variety “Otome Pink” and found that petiole length was shorter with increasing dose of mutagenic agents. Mahure et al.2010 recorded minor dosages like 10, and 20 Gy enlarged leaf area, but 30 Gy dwindled leaf area over control. In yet another study by Dilta et al. 2006 a reduction in leaf number was reported in Dendranthemum grandiflorum kitam cv. “Gulmohar” under gamma irradiation dose range of 1.0–3.0 kR.

Influence of gamma irradiation on days to floral bud appearance and plant height (cm) at flowering

With the increment of each dose of irradiation (Table 4), there was a considerable delay in days to bud appearance contrast to control plants (23.50). Under 10, 20, and 30 Gy doses, days to bud appearance were recorded 27.25, 37.00, and 39.25, respectively. Whereas, days to bud appearance under the last dose of 40 Gy were recorded significantly highest 40.75, representing maximum delay compared to control (Fig. 3i). The results in the present study may be due to the disturbances in the biochemical pathway, which assists in the synthesis of flower-inducing substances and hence delay in flowering.

Table 4

Influence of ⁶⁰Co gamma irradiation on the number of days to floral bud appearance and plant height at flowering in Chrysanthemum (*Dendranthemum morifolium* L.) cv. “Candid”
Dose	Number of days to floral bud appearance	Plant height at flowering(cm)
0 Gy	23.50a ± 1.73	53.25a ± 2.50
10 Gy	27.25b ± 1.50 (15.95)	49.00b ± 0.82 (7.98)
20 Gy	37.00 c ± 0.82 (57.44)	36.50c ± 0.58 (31.45)
30 Gy	39.25cd ± 1.89 (67.02)	34.00d ± 20.82 (36.15)
40 Gy	40.75d ± 2.22 (73.40)	31.50e ± 1.00 (40.84)
L.S.D _P≤0.05	2.58	2.03
Figures in the parenthesis represent per cent increase in case of days to floral bud appearance and per cent decrease in plant height at flowering over control

The present study results concur with the findings of Datta and Banerji 1993 who observed delayed flowering behaviour after irradiating rooted cutting of small decorative type chrysanthemum cv. “Kalyani Mauve.” In another study Dilta et al. 2006 also observed a significant delay in days to bud formation, buds showing colour, and days for full bloom in the treated plants, often chrysanthemum cultivar as compared to control. Similar were the results recorded by Misra et al. 2009 in chrysanthemum cultivar “Pooja.” Plant height is the genetic characteristics of the plants, and it was expressed as per the individual potential of a variety. With the increment of each dose of irradiation (Table 4), there was a major decline in plant height compared to control plants (53.25 cm). Under 10, 20, and 30 Gy doses, plant height was recorded at 49.00, 36.50, and 34.00 cm, respectively. Whereas, plant height at the flowering time under the last dose of 40 Gy was recorded significantly lowest 31.50 cm, representing the highest decrease compared to control (Fig. 3j). Reduction in vegetative characters by gamma rays treated plants depends on the nature and degree of chromosomal injury or morphological, cytological and physiological, disturbance induced by irradiation and the decline of interior auxin manufacture, leading to plummeting growth of the plant (Banerji and Datta 2002). Also, due to the inactivation and decrease in auxin synthesis and nature and degree of chromosomal aberration (Singh et al. 2011). The results in the present study are in concurrence with the findings of Banerji and Datta 1993. They observed a decrease in plant height with an increased dose of gamma irradiation in the rooted cutting of chrysanthemum cv. “Jaya” and “Lalima.” Kole and Meher 2005 also reported decreasing effects in zinnia might be due to physiological damage caused by mutagen at higher doses.

Influence of mutation frequency and gamma irradiation on flower colour

With response to the colour of flowers after irradiation, desired colour mutants were selected only from the plants irradiated with 10 Gy dose, which evolved 60 per cent of pink, 15 per cent of orange-pink, 10 per cent white, 5 per cent light yellow (5%) and remaining 10 per cent were as identical as control, i.e., showing original red colour (Fig. 3-a, b, c, and d). Higher doses of 20, 30, or 40 Gy produced either distorted red buds or distorted red (Fig. 3-e and f). Colour mutants under 20, 30, and 40 Gy were undesirable. The results in the present study may be due to physiological transformations which happen in the plant; hence, deferred flowering occurs at elevated doses due to inhibitory result. This can be ascribed to the fact that no chimeric growth was developed in the shoot due to mutagenesis. Tissue or shoot without chimeric growth leads to non-formation, diverse colour variations in petals as testified in chrysanthemum by Longton 1980. This quoted observation is in close conformity to the present study. Data regarding the mutation frequency in chrysanthemum flowers based on flower colour showed a highly desired mutation frequency amounting to 90 per cent when the plants were irradiated with 10 Gy dose. Whereas, under 20, 30, and 40 Gy doses flower mutation frequency, although recorded cent per cent, produced undesirable mutants. The results recorded in the present study are in accordance with the finding of Siavash et al.2009 who advocated increased mutation frequency when plants were UV irradiated.

The investigation concludes that irradiation 40 Gy dose resulted in a significant decrease in per cent rooting, subsequent decline in field survival, days to floral bud appearance, plant height at flowering, and mutation frequency. The highest number of desired mutants concerning flower colour (Light pink, Orange- Pink, White, and yellow) and the highest mutation frequency were observed in shoots irradiated with 10 Gy. Hence, 10 Gy gamma irradiation treatment is congenial for mutagenesis in Chrysanthemum Cv. “Candid.”

GA₃ : Gibberelic acid: MS : Murashige; Skoog’s medium BAP: 6-Benzyl amino purine;; Gy : Grays; PANBIT: Panoramic Batch Irradiator

Acknowledgements

We want to express our sincere gratitude to UGC, GoI for the MANF fellowship.

Conflict of interest

No conflict of interest.

Declarations

Not applicable

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Congenial in Vitro γ-Ray Induced Mutagenesis Underlying the Varied Array of Petal Colours in Chrysanthemum (Dendranthemum Grandiflorum Kitam). ‘Candid’

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Abstract

Figures

Introduction

Materials And Methods

Results And Discussion

Influence of γ- rays on leaf area and number of leaves

Conclusion

Abbreviations

Declarations

References

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