Rapid micropropagation and callus induction of Terminalia bellerica Roxb. - An endangered plant

Similar documents
Clonal micropropagation and callus induction of Terminalia bellerica- An endangered plant

An efficient protocol for clonal micropropagation of Mentha piperita L. (Pipperment)

Plantlet Regeneration via Somatic Embryogenesis in Four Species of Crocus

SOMATIC EMBRYOGENESIS OF DREPANOSTACHYUM FALCATUM AN IMPORTANT HILL BAMBOO-A RAPID MEANS OF MICROPROPAGATION

Summary and conclusion

Keywords: Agarwood, satalum, resin, micropropagation, tok

Effect of Cytokinins on Multiple Shoot Regeneration from Leaf Derived Callus of Inula

Plant Regeneration via Organogenesis and Somatic Embryogenesis

MICROPROPAGATION OF JATROPHA CURCAS (L.)

International Journal of Pharma and Bio Sciences RAPID IN VITRO PROPAGATION TECHNIQUE FOR SUGARCANE VARIETY 018

In vitro propagation of Musa sp (Banana)

CHAPTER 2. IN VITRO REGENERATION OF Gerbera jamesonii Bolus Ex. Hook f. Previous research has proven that Gerbera jamesonii could successfully be

Gregor Mendel Foundation Proceedings 2007:

MICROPROPAGATION OF RICE (ORYZA SATIVA L. CV SWAT-II) THROUGH SOMATIC EMBRYOGENESIS

International Journal of Pharma and Bio Sciences DIRECT REGENERATION FROM EMBRYO CULTURES OF LYCOPERSICON ESCULENTUM MILL CV PUSA RUBY ABSTRACT

Low-Cost Alternatives for Conventional Tissue Culture Media

MICROPROPAGATION OF CHRYSANTHEMUM (CHRYSANTHEMUM MORIFOLIUM) USING SHOOT TIP AS EXPLANT

Available online at

Micropropagation of Chlorophytum borivilliens through direct organogenesis

In vitro Plant Regeneration of Withania somnifera. Ujjwala Supe, Fanisha Dhote and M.G. Roymon

Environmental and Genotypic Effects on the Growth Rate. of in Vitro Cassava Plantlet

In vitro regeneration performance of Corchorus olitorius

In Vitro Microcorm Formation in Saffron (Crocus sativus L.)

INFLUENCE OF PLANT GROWTH REGULATORS ON IN VITRO CLONAL PROPAGATION OF DENDROBIUM SONIA EARSAKUL

Direct Regeneration of Shoot from Axillary Bud of Citrus Reticulate

Micro propagation of sugarcane (Saccharum officinarum L.) through auxiliary buds

IN VITRO GROWTH AND DEVELOPMENT OF DENDROBIUM HYBRID ORCHID. H. KHATUN 1, M. M. KHATUN 2, M. S. BISWAS 3 M. R. KABIR 4 AND M. AL-AMIN 5 Abstract

Received : Accepted:

Impact of Monochromatic Lights Treatment on In Vitro Regeneration of Celastrus paniculatus

Micropropagation of Salvia broussonetii Benth. - A Medicinal Plant Species

Regeneration of Asparagus racemosus by shoot apex and nodal explants

IN VITRO MASS MULTIPLICATION OF NONI (Morinda citrifolia L.) THROUGH NODAL SEGMENT EXPLANTS

SOMATIC EMBRYOGENESIS AND REGENERATION OF PLANTLET IN SAFFRON, CROCUS SATIVUS L.

In Vitro Regeneration of Aloe Vera (Aloe barbadensis Mill)

In Vitro Flowering and Shoot Multiplication from Nodal Explants of Ceropegia bulbosa Roxb. var. bulbosa

Researcher, 1(3), 2009, Micropropagation Of Prosopis Cineraria (L.) Druce A Multipurpose Desert Tree

EFFECTIVE CALLUS INDUCTION AND PLANT REGENERATION IN BRASSICA NAPUS (L.) VAR DGS-1

EFFECT OF BENZYLAMINO PURINE AND NAPHTHALENE ACETIC ACID ON CALLUS AND PROTOCORM FORMATION OF DENDROBIUM CV. BANYAT PINK

In vitro REGENERATION OF MUNGBEAN (Vigna radiata L.) FROM DIFFERENT EXPLANTS

Int.J.Curr.Res.Aca.Rev.2017; 5(8): 12-17

MULTIPLE SHOOT REGENERATION IN DENDROBIUM FIMBRIATUM HOOK AN ORNAMENTAL ORCHID ABSTRACT

IN VITRO PLANT REGENERATION STUDIES IN BRINJAL

AVOCADO CALLUS AND BUD CULTURE

In vitro Clonal Propagation of Vitex negundo L. An Important Medicinal Plant. M.R. Islam, Ruseli Khan, S.N. Hossain, G. Ahmed and L.

IN VITRO SHOOT MULTIPLICATION AND CALLUS INDUCTION IN GLADIOLUS HYBRIDUS HORT.

The effects of BA and NAA on multiplication of Butterwort (Pinguicula gigantea) in vitro

Regeneration potential of seedling explants of chilli (Capsicum annuum)

In vitro Conservation of Rose Coloured Leadwort: Effect of Mannitol on Growth of Plantlets

Callus induction and plant regeneration from leaf explants of Spilanthes acmella Murr. : An endangered medicinal plant

Researches regarding bitter melon (Momordica charantia) in vitro regeneration

In vitro Regeneration of Geranium (Pelargonium graveolens L. Hert.) through Axillary bud Culture an Important Essential oil yielding plant

American-Eurasian Journal of Sustainable Agriculture, 6(4): , 2012 ISSN Somatic Embryogenesis and Plantlet Regeneration in Amla

Seed Culture of Aromatic Rice Varieties Under Salt Stress

A micropropagation system for Eucalyptus dunnii Eucalyptus sp

ORGANOGENESIS IN CHRYSANTHEMUM MORIFOLIUM RAMAT (CULTIVAR ROMICA ) CALLUS CULTURES SMARANDA VÂNTU

In vitro callus induction, regeneration and micropropagation of Solanum lycopersicum

Investigation of Plant Growth Regulators Effects on Callus Induction and Shoot Regeneration of Bunium persicum (Boiss.) B. Fedtsch.

In vitro clonal propagation of the neem tree (Azadirachta indica A. Juss.)

VETIVER PROPAGATION. Nurseries and Large Scale Propagation. Dr Paul Truong Veticon Consulting Brisbane, Australia

SK. Jaffar * et al. /International Journal Of Pharmacy&Technology

Plantlet regeneration of Pinus kesiya Royle ex Gord. from mature embryos

Organogenic plant regeneration via callus induction in chickpea (Cicer arietinum L.) Role of genotypes, growth regulators and explants

Comparison of Regeneration Efficiency of Three Different Genotypes of Basmati Rice under in Vitro Condition

In Vitro Regeneration of Parthenocarpic Cucumber (Cucumis sativus L.)

Rapid in vitro regeneration of Gerbera jamesonii (H. Bolus ex Hook. f.) from different explants

In Vitro Shoot Proliferation from Excised Shoot Tip and Nodal Segment of Anacardium occidentale L.

Plant regeneration through direct shoot bud formation from leaf cultures of Paphiopedilum orchids

In vitro propagation of muskmelon (Cucumis melo L.) from nodal segments, shoot tips and cotyledonary nodes

Effect of BA NAA and 2,4-D on Micropropagation of Jiaogulan (Gynostemma pentaphyllum Makino)

MICROPHOPAGATION OF DATE PALM (PHOENIX TISSUE CULTURE TECHNIQUE

4.1. Germplasm collection The state of Tamil Nadu is located in the southern eastern part of Indian

International Journal of Sustainable Crop Production (IJSCP)

Rapid Micropropagation and Callus Induction of Catharanthus roseus in Vitro Using Different Explants

H. E. Sommer, H. Y. Wetzstein and N. Lee

Full length Article Studies on callus induction and shoot regeneration in Tomato

Genotype dependent influence of phytohormone combination and subculturing on micropropagation of sugarcane varieties

Adult Plants and Juvenile Seedlings of Persimmon (Diospyros kaki L.)

Organogenic plant regeneration via callus induction in Stevia rebaudiana Bert

THE EFFECT OF DIFFERENT PLANT GROWTH REGULATORS ON SHOOT INDUCTION OF Orthosiphon aristatus Boldingh.

EFFECT OF DIFFERENT CULTURAL CONDITIONS ON MICROPROPAGATION OF ROSE (ROSA INDICA L.)

Original Papers Plant Tissue Culture Letters, 11(3), (1994) INouE**, Koji NoMuRA***, Seiko TAKAHASHi**, OsHIMA* and Kiyoshi MASUDA**

Shoot Proliferation of Dendrobium Orchid with BAP and NAA

CONSIDERATIONS REGARDING THE EFFECTS OF GROWTH REGULATORS OVER THE IN VITRO MORPHOGENETIC REACTION AT ORIGANUM VULGARE L.

Albino Regenerants Proliferation of Dendrocalamus asper in vitro

Effects of Different Concentrations of α-naphthaleneacetic Acid and 6-benzylaminopurine on Shoot Regeneration of Vinca minor L.

International Journal of Current Biotechnology

In vitro Propagation of Euphorbia pulcherrima Willd. Through Somatic Embryogenesis

In-vitro Regeneration of Bambusa balcooa (Bamboo) through Nodal Segments

Sweta Mishra and Yuvraj K Vala

In-vitro callus induction and shoot regeneration in Physalis minima L

Albino Regenerants Proliferation of Dendrocalamus Asper In vitro

Effect of BA and 2iP on Shoot Proliferation and Somaclonal Variation of Gardenia jasminoides Ellis in vitro Culture

In Vitro Mass Multiplication of Anthurium andreanum (HORT) Cultivars

Effect of Growth Regulators on In vitro Cultures of Aconitum heterophyllum: An Endangered Medicinal Plant

ESTABLISHMENT OF A SUITABLE AND REPRODUCIBLE PROTOCOL FOR IN VITRO REGENERATION OF GINGER (ZINGIBER OFFICINALE ROSCOE)

Chapter 21. Micropropagation of Cordyline terminalis. Tui Ray, Prasenjit Saha, and Satyesh C. Roy. Abstract. 1. Introduction

The application of leafy explant micropropagation protocol in enhancing the multiplication ef ciency of Alstroemeria

MASS PRODUCTION OF A RARE AND ENDANGERED SPECIES, ASTRAGALUS MEMBRANACEUS VAR. ALPINUS NAKAI, THROUGH AXILLARY BUD CULTURE AND IN VIVO ROOTING TEST

In Vitro Regeneration of Papaya (Carica papaya L.) Variety Surya

Efficient micropropagation of Vanilla planifolia Andr. under influence of thidiazuron, zeatin and coconut milk

Transcription:

Available online at www.pelagiaresearchlibrary.com Asian Journal of Plant Science and Research, 2012, 2 (3): 364368 ISSN : 22497412 CODEN (USA): AJPSKY Rapid micropropagation and callus induction of Terminalia bellerica Roxb. An endangered plant Jitendra Mehta*, Monika Sain, Banwari Lal Mathuriya, Ritu Naruka, Ambika Kavia and Dev Ratan Sharma Plant Tissue Culture Laboratory & Department of Biotechnology, Vital Biotech Research Institute, University of Kota, Kota (Rajasthan), India. ABSTRACT An in vitro micropropagation system has been developed for Terminalia bellerica Roxb., an important Indian medicinal plant. Nodal segments obtained from 15dold aseptically grown seedlings were used as explants. MS medium containing 2.0 mg/l BAP was found most suitable for culture initiation. Although shoot multiplication was achieved on MS medium containing BAP and Kn, the maximum number of shoots was obtained with 3.5 mg/l BAP+ 0.5 mg/l Kn. Best rooting response was observed on medium containing quarter strength MS salts, 0.8% agar and 1.0 mg/l IBA. Plantlets were hardened initially in culture room conditions and then transferred to misthouse. Maximum callus induction response was observed on MS medium supplemented with 0.25 mg/l 2,4D+ 0.3 mg/l Kn within 4 weeks from leaf petiole. Keywords: Medicinal plant, callus, Bahera, micropropagation, shoot multiplication. INTRODUCTION Plants containing beneficial and medicinal properties have been known and used as sources of food, fodder, oils, medicines, fuel, wood, fibers, and timber by increasing population growth. Due to increased demand for pulp, paper, construction materials, farmlands and fuel, status of woody trees especially forest trees are greatly affected (Giri et al. 2004). Plant is our natural wealth and its conservation is important for economic, ecological, scientific, medicinal and ethical reasons. Therefore, there is a great need to conserve forest ecosystems by agrotechnology. In recent years, in vitro approaches have been used as an efficient tool for micropropagation of trees and it proved that tissue culture technology is suitable for largescale propagation of trees in short time (Pena and Seguin 2001). Propagation of woody trees through tissue culture has many advantages over conventional propagation method like fast multiplication of the important genotypes, quick release of improved cultivars, production of diseasefree plants, seasonindependent production of plants, germplasm conservation and facilitating their easy exchange (Asthana et al. 2011). Terminalia bellerica Roxb. (Family Combretaceae) commonly known as Bahera, is a perennial deciduous tree of the tropics and found in south Asia, including Aravalis in Rajasthan, India. The fruits possess antioxidant, antistressor and immunostimulant properties and form an important ingredient of many ayurvedic preprations, such as Triphla. Conventional methods of multiplication of T. bellerica have proved inadequate on account of hard seedcoat, heavy insect infestation of seeds and low survival rate of cuttings and, therefore, require alternative methods of propagation. In recent years, plant tissue culture techniques have been employed for multiplication of various tree species using seedling and mature explants. Micropropagation of T. bellerica using mature node has 364

Jitendra Mehta et al Asian J. Plant Sci. Res., 2012, 2 (3):364368 been earlier reported by Roy et al, (1987) while Bilochi was able to establish shoot cultures using seedling nodes. However, the efficiency of multiplication was reported significantly low. Present investigation was carried out to develop an efficient protocol for micropropagation of T. bellerica from nodal explants of seedling. MATERIALS AND METHODS The fruits of T. bellerica Roxb. were collected from garden of Ummed Bhawan Place Kota, Rajasthan. They were dried and seeds were taken out after breaking the hard seedcoat. The seeds were surface sterilized subsequently with running tap water, labolin soap solution and finally with 0.1% HgCl 2 for 15 min under laminar bench. After rinsing for 56 times with autoclaved distilled water, they were inoculated aseptically on to water agar (0.8%) for germination. Cotyledonary and epicotyledonary nodes obtained from 15dold seedlings were implanted vertically on to different culture media [Murashige and Skoog (MS)] containing 2.0 mg/l BAP. Different concentrations of BAP and Kn (1.05.0 mg/l) were used individually for proliferation of shoots from seedling nodes. Combination of BAP and Kn were used for shoot proliferation. Same experiments were repeated for shoot multiplication. The medium containing 3% sucrose was solidified with 0.8% agar (Qualigens). The ph of the media was adjusted to 5.9+0.02 with 1 N NaOH or 1 N HCl solutions prior to autoclaving. Media poured in culture vessels were steam sterilized by autoclaving at 15 lbs for 1520 min. The cultures were incubated under controlled conditions of temperature (25±2 C), light (2000 2500 lux for 16 h/d provided by fluorescent tubes) and 6070% humidity. For each experiment a minimum of 7 replicates were taken and experiments were repeated thrice. Observations were recorded after an interval of 3 wk. Once culture conditions for shoot induction from explants were established, the shoots produced in vitro were sub cultured on fresh medium every 3 wk. Different combination of Auxins (0.25 mg/l) (2, 4D) and Cytokinine (BAP and Kn) (0.10.3 mg/l) were used for callus induction from leaf petiole. Rooting of elongated shoots was attempted under in vitro conditions. Auxins (IBA) alone in different concentrations (0.52.5mg/l) were incorporated in the agar (0.8%) solidified medium containing 1/4 MS salts and 1.0% sucrose. The in vitrorooted plantlets were transferred to culture bottles 1/4th filled with Soilrite composition (soil: sand: peat moss) and irrigated with 1/4 MS salt solution. These bottles were kept in controlled environmental conditions of culture room. After 3 wk of growth, the plantlets were transferred to misthouse for further growth. RESULTS AND DISCUSSION The epicotyledonary and cotyledonary (after removal of cotyledons) nodes when inoculated on MS medium containing BAP and Kn in the range 1.05.0 mg/l showed enhanced shoot proliferation. BAP at its 2.0mg/l concentration evoked best response. Incorporation of NAA or IAA improved bud proliferation but the shoots remained stunted. Shoots after their initial proliferation on medium containing 20mg/l BAP were subcultured on same fresh medium after every 21 days. Incorporation of BAP or Kn into MS medium supported multiplication of shoots in culture, BAP proved to be a better choice than Kn and the maximum number of shoot was obtained on its 2.0 mg/l concentration (Table 1, Fig. AB). When BAP was used in combination with Kn (0.5mg/l), a variety of responses were observed (Table 2, Fig. C). But best response was observed on medium containing 3.5 mg/l BAP + 0.5 mg/l Kn (Average number of shoots 5.02±0.76) and best shoot length was observed on medium containing 2.0 mg/l BAP+ 0.5 mg/l Kn (Average shoot length 3.87±0.39 cm). The full or half strength of MS medium without any PGR was failed to induce rooting of regenerated shoots. However, shoots were capable to induce root when cultured on medium containing auxins. Auxins in different concentration induced rooting when incorporated in the medium containing ¼ of MS salts. The best rooting response, however, was observed on medium containing 1.0mg/l IBA, where roots measuring 1.68±0.32 cm (average) were formed (Table 3, Fig. D). In vitro rooted plantlets were initially hardened in culture room conditions where leaves expanded. After 3 week, the plantlets were shifted to mist house. There was an increase in length of shoots and new leaves emerged which expanded quickly (Fig. EF). Leaf petiole explants were used for the purpose of callus induction. Highest diameter of callus was observed on MS Medium fortified with 0.25 mg/l 2,4D+ 0.1 mg/l BAP (callus diameter 3.5 cm) and 0.25 mg/l 2,4D+ 0.3mg/l Kn (callus diameter 3.8 cm), (Table 4, Fig. GH). 365

Jitendra Mehta et al Asian J. Plant Sci. Res., 2012, 2 (3):364368 Fig. 1 (AH) Clonal micropropagation of T.bellrica from nodal shoot explant Table 1. Effect of Cytokinin (BAP and Kn) on shoot proliferation from nodal shoot explant of T.bellrica BAP 1.0 2.0 3.0 4.0 5.0 Response (%) Kn 30 80 60 55 40 No. of Shoot/explant (mean±sd) 0.80±0.41 3.80±0.77 2.40±0.51 1.00±0.51 0.50±0.07 Shoot length (in cm) (mean±sd) 0.52±0.04 2.04±0.59 1.38±0.12 0.88±0.10 0.82±0.70 1.0 2.0 3.0 4.0 5.0 0 20 35 55 60 0.60±0.50 0.96±0.18 1.26±0.32 1.54±0.31 0.70±0.25 0.80±0.28 1.38±0.98 1.82±0.27 Means having the same letter in each Colum, do not different significantly at P< 0.05 (Tukey s test) 366

Jitendra Mehta et al Asian J. Plant Sci. Res., 2012, 2 (3):364368 Table 2. Interactive effect of Cytokinin (BAP+ Kn) on shoot multiplication by sub culture of shoot clumps of T.bellrica Shooting No. of Shoot/explant Shoot length (in cm) Response (%) 0.5 BAP + 0.5 Kn 1.19±0.21 1.44±0.80 64 1.0 BAP + 0.5 Kn 1.52±0.15 1.70±0.14 67 1.5 BAP + 0.5 Kn 2.26±0.24 2.60±0.51 74 2.0 BAP + 0.5 Kn 2.31±0.48 3.87±0.39 76 2.5 BAP + 0.5 Kn 3.78±0.57 3.06±0.22 80 3.0 BAP + 0.5 Kn 4..98±0.74 2.84±0.98 87 3.5 BAP + 0.5 Kn 5.02±0.76 2.17±0.47 90 Means having the same letter in each Colum, do not different significantly at P< 0.05 (Tukey s test) Table 3. Effect of Auxin (IBA) on root induction from isolated shoot of T.bellrica Rooting No. of roots/explants Root length (in cm) Response (%) 0.5 IBA 2.80±0.73 0.43±0.33 68 1.0 IBA 3.60±0.51 1.68±0.32 90 1.5 IBA 1.40±0.52 1.06±0.08 82 2.0 IBA 1.38±0.37 0.92±0.10 78 2.5 IBA 1.08±0.19 0.51±0.05 73 Means having the same letter in each Colum do not different significantly at P< 0.05 (Tukey s test) Table 4. Effect of different Hormones on Callus proliferation and Morphology Hormone Conc. (mg/l) Callus diameter after 7 weeks subculture (cm) 2,4D + Kn 3.2 2.6 3.8 0.25 + 0.1 0.25 + 0.2 0.25 + 0.3 2,4D + BAP 0.25 + 0.1 0.25 + 0.2 0.25 + 0.3 3.5 2.8 2.4 Callus proliferation Scoring +++ ++ ++++ ++++ ++ +++ ++++ Intense +++ Moderate ++ Meager CONCLUSION Color of callus Green Whitish green Dark green Dark green Whitish green Green Morphology of callus Compact Friable Nodular Nodular Friable Compact The seedling derived explants, being juvenile, are frequently used for micropropagation, as they are easy to establish in culture. In T. bellerica, MS medium containing 2.0mg/l BAP was the best for culture initiation. We have found that T. bellerica culture grew better on MS medium in comparison to other media. In T. bellerica, 2.0 mg/l BAP was most suitable for shoot multiplication. We also observed improvement in shoot multiplication by different concentrations of BAP (0.53.5 mg/l) in medium along with Kn (0.5 mg/l). Best shooting response was observed on media containing 3.5 mg/l BAP+ 0.5 mg/l Kn (Average number of shoots 5.02) and 2 mg/l BAP+ 0.5 mg/l Kn (Average shoot length 3.87 cm). IBA (Auxin) has been widely used as root induction hormone under in vitro and in vivo condition. We also found positive role of IBA during in vitro rooting. In T. bellerica, 1.0mg/l IBA proved to be the best for in vitro rooting. The in vitro rooted plants were hardened first under controlled conditions of culture room and then shifted to mist house where they exhibited and hence, growth and 90% survival. Most responsive callus induction was observed on MS Medium supplemented with 0.25 mg/l 2,4D+ 0.1 mg/l BAP. Acknowledgement We are grateful to Plant Tissue Culture Division, Vital Biotech Research Institute, Kota for providing laboratory facilities and also thankful to Mr. Jitendra Mehta of Vital Biotech for sincere efforts in writing this research paper. 367

Jitendra Mehta et al Asian J. Plant Sci. Res., 2012, 2 (3):364368 We are also grateful to Mrs. Monika, Mr. Banwari Lal Mathuriya, Miss Ritu Naruka, Miss Ambika Kavia, and Mr. Dev Ratan Sharma for their continuous team work. We would also like to thank Mrs. Jaya Mehta and staff members of Vital Biotech Research Institute for encouragement and we would like to thank the reviewers of this paper for their excellent comments. REFERENCES [1] Giri C, Shyamkumar B, Anjaneyulu C, Trees Struct Funct, 2004, 18:115 135. [2] Pena L, Seguin A, Trends Biotechnol, 2001, 19:500 506. [3] Asthana P, Jaiswal VS, Jaiswal U, Acta Physiol Plant, 2011, 33:1821 1829. [4] Roy S K, Pal P K & Das A K, Bangladesh J Bot, 1987, 16: 125130. [5] Bilochi G, In vitro studies on some medicinal plants of Aravallis in Rajasthan. PhD Thesis, Mohanlal Sukhadia University, Udaipur, India, 2002. [6] Purohit S D & Singhvi A, Sci Hortic, 1998, 76: 219229. [7] Minocha S C, Plant growth regulators and morphogenesis in cell and tissue culture of forest trees, in Cell and tissue culture in forestry, General principles and biotechnology, edited by J M Bonga and D J Durzan (Martinus Nijhoff Publishers. Dodrecht, The Netherlands), 1987, 1: 5256. [8] Gamborg O L, Miller R A & Ojima K, Exp Cell Res, 1968, 50: 151158. [9] Feyissa T, Welander M. & Negash L, Plant Cell Tissue Organ Cult, 2005, 80: 119127. [10] White P R (Ed), the cultivation of animal and plant cells (Ronald Press, New York) 1963. [11] Thorpe T A, Morphogenesis and regeneration, in Plant cell and tissue culture, edited by I K Vasil and T A Thorpe (Kluwer Academic Publishers, The Netherlands) 1994, 1736. [12] Murashige T & Skoog F, Physiol Plant, 1962, 15: 473497. [13] AitkenChristie J & Connett M, Micropropagation of forest trees, in Transplant production systems, edited by K Kurata and T Kozai (Kluwer Academic Publishers, The Netherlands) 1992, 163194. [14] Schenk R V & Hildebrandt A C, Can J Bot, 1972, 50: 199204. [15] Llyod G & McCown B, Int Plant Prop Soc Comb Proc, 1980, 30: 421427. [16] Kretzschmar U, Silvae Genet, 1993, 42: 163168. 368