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The effect of gibberellic acid and benzyladenine in growth and flowering of Lily (Lilium longiflorum).

Introduction

Lily flower (Lilium longiflorum Thunb.) is commercially produced as a potted plant in the US. The most important factor determining its market value is the accuracy and uniformity with which the crop flowers in time for sales. Gibberellins and cytokinin are plant growth regulators that affect many phases of plant growth and development from germination, leaf expansion and shoot elongation to flowering, fruit development and mobilization of seed reserves. Potential of using growth regulators specially Gibberellins and cytokinin is not studied in Lilium sufficiently.

Emongor and Tshwenyane [6] reported that cytokinins, increase chlorophyll development and chlorophyll synthase. Chlorophyll degradation in leaves of cut flowers is controlled by gibberellins [27]. Sing et al [25] said that gibberellic acid treatment cause to increase in longevity of gladiolus cut flowers. Mutuee et al [18], said that gibberellins increase fresh weight of cut flowers.

The aim of this study is to study on some important traits of lily cut flowers treated with gibberellic acid and 6-benzyladenine.

Material and method

In this study we used Lilium hybrid [Asiatic.sup.x] Longiflorum c.v. Menorca. 108 forced bulbs with 16-18 centimeter diameter were supplied from Khadem greenhouse in Pakdasht, Varamin, Iran. I.R. An experiment was performed by using factorial experiment arranged in a completely randomized design with three replications. Also, we used two factors, gibberellic acid and 6-benzyladenine, each in three levels (0, 75 and 150 ppm). Before planting lilium bulbs were disinfected with 1% (w/v) benomyl fungicide. Bulbs were in treatment for 24 hours before culture. These treatments were contents of control, 75 ppm gibberellic acid, 150 ppm gibberellic acid, 75 ppm 6-benzyladenine, 150 ppm 6-benzyladenine, 75 ppm gibberellic acid with 75 ppm 6-benzyladenine, 150 ppm gibberellic acid with 150 ppm 6-benzyladenine, 75 ppm gibberellic acid with 150 ppm 6-benzyladenine and 150 ppm gibberellic acid with 75 ppm 6-benzyladenine. In order to supply perfect temperature for lily growth, day temperature was regulated at 18-22[degrees]C and night temperature was regulated at 16-18[degrees]C [4,5]. Photoperiod was 13 hours and light density in greenhouse was 302 micro molz[m.sup.-2]x[s.sup.-1] which is measured by illuminati meter (model INS-DX-200).In this study, flowers were harvested when flower bulbs colored completely. Flower harvesting was down at morning. Cut flowers were quickly transferred to horticultural laboratory. Then cut flowers weighed and lodged in equal glasses with distinct distilled water. The cut flowers were stored at room temperature with 70% Relative humidity. Light value was 21 micro mol / [m.sup.2] / s, which was measured by illuminati meter.

Data were subjected to ANOVA procedure in SAS (ver. 9.0, SAS Institute, Inc.) and appropriate means were separated using the LS means.

Result:

Number of Yellow Leaf:

Variance analysis showed that effects of gibberellic acid and interaction effect of gibberellic acid and benzyladenine was significantly positive (p < 0.05) but effects of 6-benzyladenine was not significant. Comparison of means showed that the maximum number of yellow leaves was related to control treatment and the minimum number was related to gibberellic acid 150 ppm (table 1).

Chlorosis is an ordinary problem in lily production. Leaf yellowing in lily during production occurs as a gradually physiological problem at a greenhouse or occurs suddenly during the storage [6,13]. Flower senescence is along with leaf fading and chlorophyll decrease [13]. In one place Gibberellic acid and benzyladenine prevent from proteolysis and chlorophyll degradation [22,23,6] and on the other hand prevent from leaf senescence and yellowing by increasing in chlorophyll synthesis and chloroplast development [6].

In addition, gibberellic acid in chloroplast membrane causes to regulate photosynthesis process and increases leaf longevity [13].

Our results is similar to results of Nowak and Mynett [19] on lily, Jordi et al [14] on Alestromeria, Han [11,12] on lily, Pun et al [22] on gladiolus, Ranwala and Miller [24] on lily and Kim and Miller [15] on tulip, confirm the positive and significant role of gibberellic acid and benzyladenine on decreasing number of yellow leaf.

Number of Flower Bud:

Variance analysis showed that effects of gibberellic acid was significantly positive (p<0.01) and effects of 6-benzyladenine and interaction effect of gibberellic acid and benzyladenine was significantly positive (p<0.05). Gibberellic acid 150 ppm with benzyladenine 75 ppm showed the maximum number of flower bud and the control treatment had the least.

Gibberellic acid may cause to increase in the accessible substances in flowering initiation time. This subject can be related to flowering increase [3]. Our results like the results of Funnel et al [8] on Zantedeschia, Ali and Elkiey, [3] on Zantedeschia, confirm the positive and significant role of gibberellic acid on increasing bud number and likewise the results of Pogroszewska et al [21] on Allium karataviense, it is showed the positive and significant role of benzyladenine.

Chlorophyll a:

Variance analysis showed that effects of gibberellic acid and benzyladenine was significantly positive (p<0.01 and p<0.05 respectively). All the treatment caused to increase in chlorophyll a content except control (table 1).

Chlorophyll Content:

Variance analysis showed that effect of gibberellic acid and interaction effect of gibberellic acid and benzyladenine on total chlorophyll content were significant (p < 0.01). Comparison of means showed in table 1.

Chlorophyll degradation in leaves of cut flowers is controlled by gibberellins [27], on the other hand, cytokinins prevent leaves senescence and delay proteolysis and chlorophyll degradation [6,28]. In addition, Emongor and Tshwenyane [6] reported that cytokinins, increase chlorophyll development and chlorophyll synthase.

Our results showed the effective role of gibberellic acid and benzyladenine on preventing of chloroplast and chlorophyll degradation. which result to decreasing in leaves senescence and increasing in total chlorophyll content. Our results is similar to results of Van Doorn and Van Lieburg [27] in Alestromeria, Gan and Amasino [9], Wingler et al [28], Guo et al [10] in chrysanthemum and Emongor and Tshwenyane [6] in Lilium.

Total Anthocyanin Content:

Variance analysis of this trait showed that gibberellic acid and interaction effect of gibberellic acid and benzyladenine on total anthocyanin content were significant (p < 0.05) and (p < 0.01) respectively. Comparison of means showed that samples, which are treated by gibberellic acid 75 ppm, benzyladenine 150 ppm, and benzyladenine 75 ppm, have the highest content of anthocyanin respectively (table 1).

Anthocyanin pathway activity in petal development duration, need a set of action and reaction between environment and developmental signals. Gibberellins, sugars and light are needed for infusing anthocyanin gene transcription and for aggregating pigments in corrolla. The effects of these signals are not just for pigment biosynthesis. They also control petal cell extension and infusion of gene expression from different ways. These reports suggest a model, which in this model gibberellins, sugar and light increase the activity of transcription regulator that is control different ways for development completion [28]. Weiss [28] reported that gibberellic acid could be a promoter for starting extension and coloration of flower. This plant growth regulator is produced in anthers, which are in developmental step and then transfer in to corolla and at last relate to physiological and biochemical processes such as petal growth and pigment production. Weiss, reported that gibberellic acid could infuse some genes of essential enzymes in anthocyanin production pathway such as chalkon synthetase, chalkon isomerase, di-hydro flavenol redoctase. Kwack et al [16] reported that gibberellic acid regulate phenyl alanin amoniliyase activity which is increase anthocyanin content in plant culture. Our results like Weiss [28] and kwack et al [16] showed that using gibberellic acid and benzyladenine increased anthocyanin content compare with control treatment.

Longevity:

Variance analysis showed that gibberellic acid and interaction of gibberellic acid with benzyladenine were significantly effective on longevity (p< 0.05). Comparison of means showed that using gibberellic acid 75 ppm related to the highest longevity. In addition, after that using gibberllic acid 150 ppm with benzyladenine 75 ppm and gibberellic acid 75 ppm with benzyladenine 150 ppm caused to the highest longevity respectively (table1).

Kwack et al [16] reported that flower senescence is a process that is caused by different processes such as oxidation actions which caused by aggregation of hydrogen peroxide radical, destruction of lipids and phospholipids structures and then increasing in ions leakage which is caused by cell membrane damage. Cytocynins delay petal senescence by protecting cells and proteins. Benzyladenine can increase cut flower longevity by improving cell membrane permanency, delaying membrane lipid peroxidation in cell and decreasing in ions leakage. In addition, benzyladenine can increase hegxos sugars (sucrose and fructose) availability in cell by increasing in a- amylase and invertase enzyme activity [17]. Our results also showed the important role of gibberellic acid and benzyladenine in significant increasing in lily longevity like Marousky [17], Guo et al [10], Ranwala and Miller [24] and Singh et al [25].

Carotenoid Content:

Variance analysis showed that effect of gibberellic acid and benzyladenine on carotenoid content were significant (p < 0.01 and p < 0.05 respectively).

Our results like the results like the results of Abdel-Wahid and Sweify [2] on Beaucarnea recurvata showed the effective role gibberellic acid in increasing carotenoid content and our results like the results of Abd El-Aziz [1] in croton plant and Penner and Wiely [20] in cucumis and cucurbita showed the effective role of benzyladenine in increasing carotenoid content.

Initial Fresh Weight:

Variance analysis of this trait showed that gibberellic acid and benzyladenine effects on fresh weight was significantly positive (p< 0.05). Comparison of means of different gibberellic acid densities showed that gibberellic acid 75 ppm had the highest initial fresh weight and the control treatment had the lowest initial fresh weight (fig 1). Comparison of means for benzyladenine showed that benzyladenine 75 ppm had the highest initial fresh weight and the control treatment had the lowest (fig 2).

Emongor and Tshwenyane (2004, Mutui et al, [18] reported that gibberellins increase cut flowers fresh weight by negation of cell water potential. Also benzyladenine increase hegxos sugar (sucrose and fructose) availability by increasing in a- amylase and invertase activity in cell and osmotic potential and at last increase the fresh weight [17].

[FIGURE 1 OMITTED]

Our results like Emongor and Tshwenyane [6] and Mutui et al [18] showed that gibberellic acid had significant positive role in increasing fresh weight and in our results benzyladenine had the significant positive role in increasing initial fresh weight too.

Petal Total Soluble Solid:

Variance analysis of this trait showed that the effect gibberellic acid on petal total soluble solid was significantly positive (p<0.05). Comparison of means of different gibberellic acid densities showed that gibberellic acid 150 ppm had the highest and the control treatment had the lowest total soluble solid (fig 3).

Gibberellic acid can increase starch hydrolysis to glucose and fructose and sucrose, which is resulted of combination of these two monosaccharide, will cause to boost of cell wall. On the other hand using gibberellic acid can cause to more production and conservation of carbohydrate in stem and petal tissues [26]. Singh et al at 2008 in studying growth regulators and sucrose on post harvest physiology, membrane persistence and gladiolus cut flower vase life reported that gibberellic acid with sucrose increase gladiolus vase life via increasing lipids peroxidation and then decrease activity of lipoxigenase enzyme and improving petal cell wall substance. Our results like Singh et al [25] on gladiolus and Taheri Shiva [26] on lily showed the positive and significant effect of gibberellic acid on petal soluble solid.

[FIGURE 2 OMITTED]

Conclusion:

As we said, results of this study and some other studies confirm that gibberellic acid and 6-benzyladenine increase some process such as cell wall tension and thus cell water potential decline [7] and more water absorption to cell and at last increase fresh weight and flower succulence [18,6]. On the other hand effect of gibberellic acid and 6-benzyladenine on increasing of carbohydrate hydrolysis induction cause to stability of respiration and thus increase of cut flowers vase life [5].

[FIGURE 3 OMITTED]

Effect of gibberellic acid and 6-benzyladenine on preventing of senescence [22,24,6] and its effect on chlorophyll synthesis and chloroplast development increasing prevent from leaves yellowing [10,6].

Cut flower harvesting and recutting them cause to stress is occurred. This stress decrease photosynthesis by effecting on tylacoeid membrane, tylacoeid osmotic potential and increase production of ROS. Carotenoids have an antioxidant role and prevent or decrease efficacies of ROS stress. On the other hand, anthocyanins prevent cell oxidation process and retard senescence that is occurred because of hydrogen peroxide aggregation and degradation of cell membrane, proteins, lipids and increase of ions leakage [28,26]. Therefore, according to result of this study and some other studies showed that increasing in anthocyanin and carotenoid would increase lily vase life. Thus, gibberellic acid interferes in anthocyanin and carotenoid biosynthesis and retard chlorophyll destroy. Therefore, a section of this effect (increase vase life) is caused to chlorophyll preservation that is attendant with leaf nitrogen level preservation.

According to these points, necessity of using growth regulator to improve quality, marketing and flower vase life is completely justification. Among treatments that we used in this study, gibberellic acid 75 ppm and gibberellic acid 150 ppm with 6-benzyladenine 75 ppm showed the high longevity and their means did not have significant difference with each other, but they showed significant difference with other treatments. Gibberellic acid 150 ppm with 6-benzyladenine 75 ppm had the most flower bud number that is showed significant difference with other treatments. Acid gibbberellic with benziladenine 75 ppm was the most effective treatment on chlorophyll a and total chlorophyll. This treatment was significantly difference with control respecting to chlorophyll a and was significantly different with control and gibberellic acid 150 ppm with 6-benzyladenine 75 ppm respecting to total chlorophyll. Gibberellic acid 75 ppm was the most effective treatment for vase life. This treatment only had significant difference with control and 6-benzyladenine 150 ppm. Gibberellic acid 75 ppm and 6-benzyladenine 75 ppm showed maximum positive effect on initial fresh weight. Gibberellic acid 75 ppm was significantly different with other treatment but 6-benzyladenine was only significantly differed from control. Gibberellic acid 75 ppm was the most effective treatment respect to carotenoid content and anthocyanin. This treatment had no significant difference with other treatment except gibberellic acid 150 ppm with 6-benzyladenine 75 ppm but gibberellic acid 75 ppm was significantly differ from all other treatment respect to anthocyanin.

Therefore according to all results and correlation of traits and in order to achieve economic aims, lily cut flowers with high quality, marketing and vase life, using gibberellic acid 75 ppm with 6-benzyladenine 75 ppm is suggested.

References

[1.] Abd El-Aziz, N.G., 2007. Stimulatory effect of NPK fertilizer and benzyladenine on growth and chemical constituents of Codiaem variegatum L. plant. American- Eurasian. J. Agric. And Environ. Sci., 2(6): 711-719.

[2.] Abdel- Wahid, S.M.K. and S.G. Sweify, 2009. Enhancement of Beaucarnea recurvata LEM.Growth by some growth regulators. Bull. Fac. Agric. Cairo Univ., 60: 2.

[3.] Ali, Y.S. and T. Elkiey, 1995. Effect of chlormequat and GA3 on growth and flowering of Calla (Zantedeschia rehmanii). J. King Saud Univ., 7(2): 271-282.

[4.] Armitage, A.M. and J.M. Laushman, 2003. Specialty cut flowers (The production of annuals, perennials, bulb and woody plant for fresh and dried cut flowers). Second edition, pp: 587- 590.

[5.] De Hortogh, A., 1996. Marketing and research requirements for Lilium in North America. ActaHorticulturae, 414: 17-24.

[6.] Emongor, V. and S.O. Tshwenyane, 2004. Effect of accel on the postharvest vase life of Easter lily. Agric. Sci., 3: 170-174.

[7.] Fathi, Gh, and B. Esmaeelpoor, 1999. Plant growth regulators. Jahade daneshgahi Mashhad, pp: 288.

[8.] Funnel, K.A., B.R. Mackay and C.R.O. Lawoko, 1992. Comparative effects of promalin and [GA.sub.3] on flowering and development of Zantedeschia "GalaXY". Acta Hort., 292: 173-180.

[9.] Gan, S. and R.M. Amasino, 1997. Making sense of senescence: molecular genetic regulation and manipulation of leaf senescence. Plant Physiol., 113: 313-319.

[10.] Guo, W., L. Zheng, Z. Zheng and W. Zheng, 2003. Phytohormones regulate senescence of cut chrysanthemum. Acta Hort., 624: 349-355.

[11.] Han, S.S., 1995. Growth regulators delay foliar chlorosis of Easter Lily leaves. Journal of American Society for Horticultural science, 120: 134-361.

[12.] Han, S.S., 1996. Enhancement of postproduction quality of Easter Lily by Growth regulators. HortScience, 31: 911-1060.

[13.] Hassanpour Asil, M., 2008. Effects of growing medium on some postharvest characteristics of cut flowers in six cultivars of Asiatic hybrid lilies (Lilium longiflorum L.). Journal of Food, Agriculture and Environment, 6(2): 442-446.

[14.] Jordi, W., G.M. Stoopen, K. Kelepouris and W.M. Van-der-Krieken, 1995. Gibberellin-induced delay of leaf senescence of Alstroemeria cut flowering stems are not caused by an increase in the endogenous cytokinin content. J. Plant Growth Regul., 14. 121-127.

[15.] Kim, H.J., and W.B. Miller, 2009. [GA.sub.4+7] plus BA enhances postproduction quality in pot tulips. Postharvest biology and technology, 51: 272-277.

[16.] Kwack, H., J. Lee and J.S. Lee, 1997. Effects of uniconazole and gibberellin on leaf variegation of ornamental plants under different light conditions. Horticultural Science, 38: 754-760.

[17.] Marousky, F.J., 1974. Shipping sulphate a new preservative for cut flower. Indian J. Hort., 31: 95-96.

[18.] Mutui, T.M., V.E. Emongor and M.J. Hutchinson, 2001. Effect of Accel on the vase life and postharvest quality of (Alestroemeria aurantiaca L.) cut flowers. Afric. J. Sci. Technol., 2: 82-88.

[19.] Nowak, J. and K. Mynett, 1985. The effect of growth regulators on postharvest characteristics of cut Lilium "Prima" inflorescences. Acta Hortic., 167: 109-116.

[20.] Penner, D. and L. Wiely, 2008. The influence of atrazine, benzyladenine and the embryonic axis on chlorophyll synthesis in cucumber and squash cotyledons. Physiologia Plantarum, 26(1): 148-151.

[21.] Pogroszewska, E., H. Laskowska, D. Wojciech, 2007. The effect of gibberellic acid and benzyladenine on the yield of (Allium karataviense Regel) "Ivory queen". Acta Sci. Pol., 6(1): 15-19.

[22.] Pun, U.K., R.N. Rowe, J.H. Rowarth, M.F. Barnes, C. Dawson and J.A. Oheyes, 1999. Influence of ethanol on climateric senescence in five cultivars of gladiolus. New Zealand. J. Crop. Hort. Sci., 27: 69-77.

[23.] Ranwala, A.P. and W.B. Miller, 1998. [Gibberellin.sub.4+7], benzyladenine and supplemental light improve postharvest leaf and flower quality of cold-stored "Star Gazer" hybrid lilies. Journal of American Society for Horticultural science, 123: 563-568.

[24.] Ranwala, A.P. and W.B. Miller, 2000. Preventing mechanism of [gibberellin.sub.4+7] and light on low-temperature-induced leaf senescence in Lilium cv.stargazer. Postharvest Biology and Technology, 19: 85-92.

[25.] Singh, A., J. Kumarand and P. Kumar, 2008. Effects of plant growth regulators and sucrose on post harvest physiology, membrane stability and vase life of cut spikes of gladiolus. J. Plant Growth Regul., 51: 221-229.

[26.] Taheri Shiva, N., 2008. Effect of plant growth regulators on anthocyanin biosynthesis in Lilium longiflorum. Master of Science thesis. Agriculture Faculty of Guilan University.

[27.] Van Doorn, W.G. and M.J. Van Lieburg, 1993. Interaction between the effects of phytochrome and gibberellic acid on the senescence of Alstroemeria pelegrina leaves. Journal of Physiology of Plant, 89: 182-186.

[28.] Weiss, D., 2000. Regulation of flower pigmentation and growth: multiple signaling pathways control anthocyanin synthesis in expanding petals. Physiologia Plantarum, 110(2): 152-157.

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(1) H. Emami, (1) M, Saeidnia, (2) A. Hatamzadeh, (3) D. Bakhshi, (4) E. Ghorbani

(1) M.Sc. Students, College of Agriculture, University of Guilan, Rasht

(2) Asso. Prof., University of Guilan, Rasht

(3) Assi. Prof., University of Guilan, Rasht

(4) PhD Student, College of Agriculture, University of Guilan, Rasht

Corresponding Author

Hossein Emami, Horticultural department, Agriculture factuality, University of Guilan
Table 1: Comparison of means of investigated traits.

                                 Carotenoid   Total
                                 content      chlorophyll

Control                          0.009 e      1.85 c
Gibberellic acid 75 ppm          0.111 a      3.09 ab
Gibberellic acid 150 ppm         0.62 bc      2.90 ab
Benzyladenine 75 ppm             0.22 cde     2.88 ab
Benzyladenine 150 ppm            0.16 de      2.68 ab
Gibberellic acid 75 ppm          0.35 cde     3.20 a
  Benzyladenine 75 ppm
Gibberellic acid 150 ppm         0.59 bcd     2.84 ab
  Benzyladenine 150 ppm
Gibberellic acid 75 ppm          0.6 bcd      2.85 ab
  Benzyladenine 150 ppm
Gibberellic acid 150 ppm         0.83 ab      2.05 bc
  Benzyladenine 75 ppm

                                                 Total
                                 Chlorophyll a   anthocyanin

Control                          0.85 b          0.26 d
Gibberellic acid 75 ppm          1.72 a          4.45 a
Gibberellic acid 150 ppm         1.52 a          2.46 bc
Benzyladenine 75 ppm             1.56 a          2.92 bc
Benzyladenine 150 ppm            1.45 a          3.31 b
Gibberellic acid 75 ppm          1.73 a          2.11 c
  Benzyladenine 75 ppm
Gibberellic acid 150 ppm         1.52 a          2.85 bc
  Benzyladenine 150 ppm
Gibberellic acid 75 ppm          1.55 a          2.51 bc
  Benzyladenine 150 ppm
Gibberellic acid 150 ppm         1.36 a          2.38 bc
  Benzyladenine 75 ppm

Means followed by similar in each column are not significantly
different at 5% level.

Table 1: Comparison of means of investigated traits.

                                 Longevity    Number of     Number of
                                              yellow leaf   flower bud

Control                          13.61 c      8.3 a         4.08 c
Gibberellic acid 75 ppm          15.77 a      6.75 b        4.75 bc
Gibberellic acid 150 ppm         15.11 ab     5.16 c        4.58 bc
Benzyladenine 75 ppm             15.44 ab     7.50 ab       4.41 bc
Benzyladenine 150 ppm            14.55 bc     7.16 ab       4.50 bc
Gibberellic acid 75 ppm          14.77 ab     6.97 ab       4.63 bc
  Benzyladenine 75 ppm
Gibberellic acid 150 ppm         15.22 ab     7.33 ab       4.91 b
  Benzyladenine 150 ppm
Gibberellic acid 75 ppm          15.52 ab     7.75 ab       5.00 b
  Benzyladenine 150 ppm
Gibberellic acid 150 ppm         15.66 a      7.16 ab       6.41 a
  Benzyladenine 75 ppm
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Title Annotation:Original Article
Author:Emami, H.; Saeidnia, M.; Hatamzadeh, A.; Bakhshi, D.; Ghorbani, E.
Publication:Advances in Environmental Biology
Article Type:Report
Geographic Code:7IRAN
Date:Jun 1, 2011
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