Improving Crop Productivity in Sustainable Agriculture (TUTEJA:CROP IMPROVEMENT O-BK) || Bamboo: Application of Plant Tissue Culture Techniques for Genetic Improvement of Dendrocalamus

13Bamboo: Application of Plant Tissue Culture Techniques forGenetic Improvement of Dendrocalamus strictus NeesC.K. John and V.A. Parasharami

Abstract

Bamboos are valuable natural resources important for both economy and environ-ment. Large-scale propagation of bamboos is difficult because of their peculiarflowering behavior only at very long intervals, and sterility in some species. Bam-boos are generally propagated by vegetative methods. Most of these methods posedifficulties such as large propagules, difficulty in extraction and transport, lack ofsufficient numbers for raising large-scale plantations, and extraction of propaguleslowering the productivity of parent clumps. In India, Dendrocalamus strictus haswide distribution, abundance, and economic importance. D. strictus occupies 53%of total bamboo area in India. In D. strictus, methods are available for (i) in vitropropagation from nodal bud explants from mature clumps, (ii) in vitro induction ofmicrorhizomes, and (iii) selecting NaCl stress-tolerant variants. D. strictus is a suit-able candidate for further work on developing abiotic stress-tolerant variants.

13.1Introduction

Bamboos are perennial, woody grasses, and are valuable natural resources impor-tant for both economy and environment [1]. They have more than 1500 docu-mented uses. Important traditional uses include housing, food, fuel, and rawmaterial for household utensils, agricultural implements, and handicrafts [2, 3].Bamboos have great potential for employment and income generation for ruralcommunities [4]. Bamboos are among the fastest growing plants on Earth [5]. Bam-boo is substitute for wood in pulp and paper manufacturing. Worldwide, more than2.5 billion people trade in or use bamboo [6]. Annual trade in bamboos (both rawand finished products) is estimated at more than US$ 5.0 billion. This representsonly a small proportion of total bamboo usage. Domestic use is estimated to accountfor at least 80% [4]. Modern manufacturing techniques allow the use of bamboo intimber-based industries, to provide bamboo flooring, board products, laminates, andfurniture [4]. Bamboo shoots are an important processed food product on the

Improving Crop Productivity in Sustainable Agriculture, First Edition.Edited by Narendra Tuteja, Sarvajeet Singh Gill, and Renu Tuteja.# 2012 Wiley-VCH Verlag GmbH & Co. KGaA. Published 2012 by Wiley-VCH Verlag GmbH & Co. KGaA.

j289

international market. China is the leading exporter with an annual export value ofnearly US$ 140 million [7]. Bamboo furniture is an expanding business in manycountries [4, 8]. Worldwide, domestic trade and subsistence use of bamboo are esti-mated to be worth US$ 4.5 billion per year. Global export of bamboo generatesanother US$ 2.7 billion [4, 6]. The vulnerability of some animal species is increasedby the simultaneous flowering and subsequent death of entire populations in supra-annual cycles ranging from 7 to 120 years or more. The best known among theseanimal species is the giant panda (Ailuropoda melanoleuca). The red panda (Ailurusfulgens) and the Himalayan black bear (Selenarctos thibetanus) are also heavily depen-dent on bamboo [4]. In recent years, with the looming threat of global warming,there is much interest in bamboo as a carbon sink [9, 10].Worldwide, there are about 1400 species of bamboos under 101–118 genera.

They are classified into three tribes: Bambuseae, Arundinarieae, and Olyreae.According to Ohrnberger [11], the subfamily Bambusoideae comprises both woodyand herbaceous bamboos with 1575 species altogether.Bamboos can be categorized into two broad categories: sympodial (pachymor-

phic) and monopodial (leptomorphic) based on the rhizome system [1]. The formeris clump forming and the latter spreading type. Most clump forming bamboos aretropical, and spreading type temperate. Clump formation in sympodial bamboos isbelieved to be an adaptation to growth in drought-prone tropical conditions, ascompact clumps and leaf shedding during summer help the plant in controllingwater loss and survive adverse climatic conditions.Ever increasing rate of deforestation makes search for alternative natural

resources an urgent need. Bamboos are suitable for large-scale cultivation in mar-ginal and degraded lands under agroforestry [12]. The extensive rhizome and rootsystems help in effectively controlling soil erosion and in reinforcing embark-ments, and accumulation of leaf mulch helps in conserving soil moisture. Fastergrowth and biomass production help in generating large amounts of oxygen andeffectively sequestering atmospheric carbon dioxide.

13.2Vegetative Propagation

Large-scale propagation of bamboos is difficult because of their peculiar floweringbehavior only at very long intervals, and sterility in some species. Bamboo seed isinfrequently produced and rapidly loses viability. Most woody bamboos are semel-parous: gregariously flower/seed only at the end of very long periods of vegetativegrowth and die en masse [1, 13]. Brandis [14] classified bamboos into three catego-ries on the basis of their flowering behavior: (i) species that flower annually ornearly so, (ii) species that flower gregariously and periodically, and (iii) species thatflower irregularly. Blatter [15] considered these three categories as fairly complete.A vast majority of the woody bamboos belong to the second category in which theintermast periods range between 10 and 120 years (or more). This peculiar flower-ing behavior has intrigued mankind for long and still remains as a botanical

290j 13 Bamboo: Application of Plant Tissue Culture Techniques

enigma. In general, the intermast periods of woody bamboos are thought of as spe-cies specific and rigid. However, it is not the case in many bamboos [13, 16, 17].Bamboos are generally propagated by vegetative methods. A vegetative propagule

of a bamboo plant must have shoot, root, and rhizome. Failure to develop any ofthese leads to failure of the propagule [18]. The most widely used vegetative propa-gation methods are clump division (offsets/rhizome), whole culm cuttings, layer-ing, culm segment cutting, and branch cutting [19]. Most of these methods posedifficulties such as large propagules, difficulty in extraction and transport, lack ofsufficient numbers for raising large-scale plantations, and extraction of propaguleslowering the productivity of parent clumps. In species such as Bambusa arundina-cea and Dendrocalamus strictus in which seeds are available once in a while, macro-proliferation of seedlings can be another method [20, 21].Bamboos in general are openly wind pollinated. They have physical/physiological

barriers that prevent self-pollination/favor cross-pollination [22–25]. Many bam-boos are known to be highly heterozygous [26–28]. This makes them suitable candi-dates for selection [29]. Most of the characters contributing to increased biomassproduction are expressed in the adult stage. Hence, propagation from adult clumpshas some advantages.

13.3Micropropagation

Micropropagation methods are available in many species of bamboo [30–40]. Planttissue culture work in bamboos started as early as 1969 [41]. During the past 40 years,a vast number of reports on in vitro propagation methods for a large number of spe-cies have appeared. Three tissue culture approaches of practical use for mass multi-plication of bamboos are (i) enhanced shoot multiplication by axillary/adventitiousbranching from juvenile and mature explants, followed by in vitro/ex vitro rooting ofthese shoots [42–44]; (ii) somatic embryogenesis from mature embryo (seed) andnodal bud explants from both seedling and mature culms [45–47]; and (iii) in vitroinduction of microrhizomes [48, 49].

13.4Genetic Improvement for Abiotic Stress Tolerance

Abiotic stresses, extremes of temperature (heat and cold), extremes of water availa-bility (drought and flood), and soil salinity pose serious threats to the growth andyield of crop plants. Vast areas of land in India and other countries are abiotic stressprone. To use these lands for cultivating crop plants, resistant varieties arerequired. There have been attempts to develop abiotic stress-resistant varieties. Ear-lier, this has been through selection and breeding [50–52] or through tissue cul-ture [53, 54], and in recent years through gene transfer [55–58]. Two mainapproaches being used to improve stress tolerance are (i) the exploitation of natural

13.4 Genetic Improvement for Abiotic Stress Tolerance j291

genetic variations, either through direct selection in stressful environments orthrough mapping quantitative trait loci and subsequent marker-assisted selection;and (ii) the generation of transgenic plants to introduce novel genes or to alterexpression levels of the existing genes to affect the degree of stress tolerance [59].The limiting factor in extension of biotechnology to abiotic stresses is the lack ofinformation on what are the “useful genes” – genes that would lead to better stresstolerance [56]. Transgenic approaches to developing stress-tolerant crop varietiesare based on the introgression of genes that are known to be involved instress response. The task of generating transgenic cultivars is not limited to thesuccess in the transformation process. Proper incorporation of the stress toleranceis essential. Evaluation of the transgenic plants under stress conditions and under-standing the physiological effect of the inserted genes at the whole plant level arechallenges remaining to be overcome [58]. Among plant biotechnologies, planttissue culture techniques offer easier and reliable methods of developing abioticstress-tolerant varieties [53, 54, 60, 61]. Bamboos are suitable candidates for devel-opment of abiotic stress-tolerant varieties given their usefulness in large-scale culti-vation in marginal and degraded lands under agroforestry.

13.5Dendrocalamus strictus

In India, D. strictus is one of the bamboo species having wide distribution, abun-dance, and economic importance. D. strictus is a compact clump forming bamboo.Clumps are deciduous and densely tufted (Figure 13.1). D. strictus occupies 53% oftotal bamboo area in India. It is widely distributed in semi-dry and dry zones alongplains and hilly tracts up to an altitude of 1000m and commonly cultivatedthroughout the plains and foothills. D. strictus grows under a wide temperaturerange from �5 to 45 �C. Most important use of D. strictus is as raw material inpaper manufacture. It is also used for construction, agricultural implements, musi-cal instruments, furniture, and so on.Among the economically important bamboos of India, D. strictus shows much

diversity for intermast periods [13]. Intermast periods reported from different partsranged between 8 and 47 years. Since this species also shows sporadic flowering,some of these reports may be that of sporadic flowering and may not represent theintermast periods. These reports point toward the existence of many land races ofD. strictus.In D. strictus, methods are available for in vitro propagation through

enhanced shoot multiplication from juvenile and mature explants [42, 62],somatic embryogenesis [45, 46], and in vitro induction of microrhizomes [48, 49](Table 13.1). Standard procedure for micropropagation involves in vitro multi-ple shoot production, followed by in vitro/ex vitro rooting and acclimatization(Figures 13.2 and 13.3).Nadgir et al. [42] obtained multiple shoots from seedling explants of D. strictus in

liquid MS medium supplemented with 2.22 mM BA and 5% coconut water. These


multiple shoots were rooted by treating them for 48 h in the dark in half-strengthMS medium supplemented with 0.49 mM IBA. Nadgir et al. [42] could obtain multi-ple shoots from nodal segment explants from mature clumps on semi-solid MSmedium supplemented with 2.22 mM BA, 0.92 mM Kn, and 10% coconut water.These multiple shoots were rooted in half-strength semi-solid MS medium contain-ing 0.25% activated charcoal after treatment for 96 h in the dark on half-strengthsemi-solid MS medium supplemented with 0.49 mM IBA. After this step, the plant-lets turned pale green and appeared in poor health. This could be avoided by trans-ferring the plantlets to liquid MS medium containing 0.44 mM BA and 0.46 mMKnþ 5% coconut water. Rooted plantlets from seedling explants as well as mature

Figure 13.1 Clump of D. strictus Nees.

13.5 Dendrocalamus strictus j293

Table13.1

Rep

ortson

microprop

agationprotocolson

D.strictusNees.

No.

Explan

tMedium

Respo

nse(s)

Reference

1.Se

edlin

gsMSþ0.88

mM

BAPþ5%

CM

þ2%

sucrose

(liquid

medium)

Multipleshoo

ts[42]

Invitrorootingof

multipleshoo

ts1/2MSþ0.49

mM

IBA(in48

hdark

follo

wed

bytran

sfer

tolig

ht)

Roo

ting80

%

Matureshoots

MSþ2.22

mM

BAPþ0.92

mM

Knþ10

%CM

þ2%

sucrose(sem

i-solid

medium)þ

0.25%

activated

charcoal

2.Se

edB5þ30

mM

2,4-Dþ2%

sucroseþ0.8%

agar

Callusform

ationin

10–12

days

atem

bryonalen

d;somaticem

bryogenesisin

67%

[45]

Callus(i)

B5þ0.5mM

IBAþ1mM

NAAþ2%

sucrose

(liquid

medium)

Con

versionof

somaticem

bryos;plan

tlet

form

ationin

40%

Plantle

ts1/2B5þ0.5mM

IBAþ1mM

NAAþ1%

sucrose

(liquid

medium

withfilte

rpaperraft)

Growth

ofconverted

plan

tlets

3.Sh

ootapices

2,4-Dmedium

Cream

ywhitecallu

s[63]

Subcultu

resfrom

(i)

Variedau

xin/cytok

inin

levels

Nodifferen

tiation

Subcultu

resfrom

(i)

VariedNAA/B

Acombinations

Green

,partiallyorganized,granularor

nod

ularcallu

s;it

was

laterconverted

toadventitiou

sshootsthatwererooted

readily

4.Produ

ctionof

somatic

embryos

MSþ13

.58mM

2,4-Dþ2.32

mM

Kn

Somaticem

bryos

[64]

Encapsulation

ofsomatic

embryos

Encapsulated

inside

Caalginatebeads

Germinationfrequen

cies

of96

and45

%wereachievedin

vitroan

din

soil,

respectively

Caalginatebeads

Add

itionalmineraloilcoatingof

Caalginatebeads

95–96

%plan

tletg

erminationun

derin

vivo

condition

s5.

Single-nod

estem

segm

ents

MSþ2.85

mM

IAAþ0.5mg/ladeninesulfate(Ads)

(mod

ified

medium)

Twoto

fouraxillaryshoots/explant

[65]

Inverselyoriented

axillaryshoots

4.9mM

IBAþ5.37

mM

NAAþ2.26

mM

2,4-Dþ1mg/lp

hloroglucinol

Roo

ting30

%


6.Nod

alexplan

tsfrom

seedlin

gcultu

reMSþ2.32

mM

Knþ9.0mM

2,4-Dþ

10mg/lA

dsþ3%

sucrose

Somaticem

bryosgerm

inated

(95–98

%)into

normal

plan

tsan

d95

%successfulroo

ting

[47]

Mature

zygo

ticem

bryo

(MZE)

MSþ2.32

mM

Knþ9.04

mM

2,4-Dþ

10mg/lA

dsþ3%

sucrose

Somaticem

bryosgerm

inated

(95–98

%)into

normal

plan

tsan

d95

%successfulroo

ting

Nod

alexplan

tsof

somaticem

bryo

regenerated

plan

ts

1/2MSþ1

.22mM

IBAþ0.5mg/lA

dsþ

1.44

mM

GA3þ3%

sucrose

Invitroflow

eringindu

ction

7.Se

edlin

gs1/2MSþ2.22

mM

BAþ2%

sucrose

Plantle

tsinitiated

[48]

Proliferationan

dprod

uctionof

plan

tlets

Simpleminim

almedia–grow

thregu

lators

Invitrorhizom

eform

ation80

%

8.Se

edlin

gsMS(liquid

medium)þ

0.88–8.88

mM

BAþ

2.32–4.65

mM

KnþCW

Multipleshoo

ts[66]

Axillary

buds

MS(liquid

medium)þ

0.88–8.88

mM

BAþ

2.32–4.65

mM

KnþCW

Multipleshoo

ts

Propagu

lesof

(i)and(ii)

MS(liquid

medium)þ

1.22–9.80

mM

IBAþ

2%sucrose

Rootindu

ction

9.Se

edMSþ30

mM

2,4-D

Embryogeniccallu

s[43]

Embryogeniccallu

sMSþ30

mM

2,4-Dþ5mM

Knþ2mM

IBAþ25

0mg/lP

VPþ3%

tablesugar

Secondary

embryogenesis(two-

tofivefold/5

weeks)

Embryogeniccallu

sMSþ30

mM

2,4-Dþ10

mM

BAPþ25

0mg/l

PVPþ3%

tablesugar

Secondary

embryogenesis(two-

tofivefold/5

weeks)

Somaticem

bryos

MSþ30

mM

2,4-Dþ5mM

NAAþ5mM

kinetin

þ25

0mg/lP

VPþ3%

sucroseþ

0.2%

Gelrite

Con

versionto

plan

tlets

10.

Embryogeniccallu

s0–20

0mM

NaC

lþMSþ3%

sucroseþ

13.6mM

2,4-Dþ2.32

mM

Knþ0.8%

agar

100mM

NaC

l-toleran

tcallusinitiated

[67]

Stable10

0mM

NaC

l-tolerantembryogenic

callu

s

MS(m

aintenan

cemedium)þ

3%sucroseþ

0.8%

agarþ9.04

mM

2,4-Dþ2.32

mM

Knþ

0–20

0mM

NaC

l

Differentiated

into

somaticem

bryos

Somaticem

bryos

tolerantto10

0mM

NaC

l1/2MSþ2%

sucroseþ0.1mM

NAAþ

0.49

mM

IBAþ10

0mM

NaC

l39

%of

maturesomaticem

bryostolerantto10

0mM

NaC

lgerm

inated

andconverted

into

plan

tlets


clump-derived explants were first transferred to 1: 1 sterile sand/soil mixture inpots. These plantlets were covered with glass beakers to maintain humidity andincubated at 25 �C and 16 h light and 8 h dark regimes. After new leaves emerged,these plantlets were transferred to greenhouse and then to field. The work byNadgir et al. [42] is one of the first reports of successful in vitro regeneration ofplantlets from a mature bamboo clump.Rao et al. [45] obtained plant regeneration through somatic embryogenesis of

D. strictus, by culturing seeds on B5 medium supplemented with 2,4-D. Calluscultures obtained from the embryonal end of the seeds differentiated intogreen embryoids. On transfer to a liquid B5 mediumþ sucroseþ IBAþNAA,40% of the embryoids developed into plantlets. Further development of theplantlets occurred in half-strength liquid B5 mediumþ 1% sucroseþ 0.5 mMIBAþ 1 mM NAA.Huang et al. [63] cultured shoot apices of D. strictus in 2,4-D-containing medium,

and developed a creamy white callus. Subculturing into media containing different

Figure 13.2 In vitro propagation of D. strictusNees. (a) Sprouting of mature clump-derivednodal bud explant on initiation medium.(b) Further elongation of the sprouted bud on

the same medium. (c) Further multiplicationand elongation of shoots on shootmultiplication medium. (d) Micropropagatedplantlets.


concentrations of auxins/cytokinins did not result in differentiation of shoots. Sub-culturing on media containing combinations of NAA and BA resulted in a green,partially organized, granular or nodular callus. This callus, on further culture,maintained its partially organized nature and produced adventitious shoots. Theseshoots were rooted and transplanted in soil.Mukunthakumar and Mathur [64] produced artificial seeds of D. strictus by

encapsulating somatic embryos obtained on MS medium containing 13.58 mM2,4-D and 2.32 mM Kn, in calcium alginate beads. Germination frequencies of 96and 45% were obtained in vitro and in soil, respectively, for these artificial seeds.The germination frequency in vivo was increased to 56% when an additional coat-ing of mineral oil on the alginate beads was tried. Germinated artificial seeds couldbe raised into plantlets.Chaturvedi et al. [65] cultured single-node stem segments excised from newly

regenerated branches of approximately 10-year-old field-grown culms of D. strictusin a modified MS medium supplemented with 2.85 mM IAA and 81.43 mM Ads.This resulted in the production of two to four axillary shoots per explant. Theseshoots did not survive because no roots were produced. Rooting was induced whenthe explants were cultured in an inverted position, in the rooting medium contain-ing 4.9mM IBAþ 5.37mMNAAþ 2.26 mM 2,4-Dþ 1mg/l phloroglucinol at pH 5.2.A seasonal effect of response was noticed by the authors, with nodal explants col-lected during July–August responding most favorably. The in vitro raised plantletswere transplanted with about 80% success. Transplanted plantlets grew normallyin soil under glasshouse as well as field conditions.Rout and Das [47] obtained plant regeneration via somatic embryogenesis in cal-

lus cultures derived from nodal explants of in vitro grown seedlings and excisedmature zygotic embryos of three bamboo species (Bambusa vulgaris, Dendrocalamus

Figure 13.3 Field trial of micropropagated and hardened plants of D. strictus Nees.


giganteus, and D. strictus) on MS medium supplemented with 2.32mM Kn, 9.5 mM2,4-D, 54.29mM Ads, and 3% sucrose incubated either in the light or in the dark.Almost all (95–98%) somatic embryos germinated into normal plants and whentransferred to soil 95% of the plants survived.Shirgurkar et al. [48] developed a micropropagation technique for the large-scale

production of D. strictus plantlets. Seedling cultures were initiated on half-strengthMS medium supplemented with 2.22mM BA and 2% sucrose. Further proliferationand production of plantlets occurred on MS mediumþ 2% sucrose. In about 80%of cultures, rhizome formation also occurred. Nearly 90% of these plantlets sur-vived when transferred to 1: 1 sand/soil mixture in pots.Ravikumar et al. [66] could induce multiple shoots from seedlings and axillary

buds of mature plants of D. strictus on MS medium supplemented with BA andKn. In the primary culture, in a span of 20–25 days about 35–45 shoots wereobtained from a nodal explant of seedling and 3–8 shoots from a nodal explantof mature plants. These multiple shoots were rooted under in vitro and ex vitroconditions. By ex vitro method using 1080 mM IBA 85–90% rooting wasachieved, when the shoots were kept at 85–90% humidity and 27–30 �C. Theshoots required 20–25 days for root initiation. Rooted plantlets performed wellduring acclimatization.Saxena and Dhawan [43] developed a complete protocol for large-scale propaga-

tion of D. strictus by somatic embryogenesis. Seeds cultured on semi-solid MSmediumþ 30 mM 2,4-D produced embryogenic callus from the embryo. Somaticembryos formed in vitromultiplied rapidly (two- to fivefold every 5 weeks) on semi-solid MS mediumþ 10 mM 2,4-Dþ 2mM IBAþ 5 mM Knþ 250mg/l soluble PVP,or MS mediumþ 10 mM 2,4-Dþ 10mM Knþ 250mg/l soluble PVP. Upon transferto MS mediumþ 5 mM NAAþ 5 mM Knþ 250mg/l soluble PVP, the dark greenembryos developed into healthy plantlets. Unrooted shoots were rooted on half-strength MS mediumþ 3 mM NAAþ 2.5 mM IBA. When these rooted plantletswere transferred to soil in polythene bags, more than 80% of them survived. Usingthis method, Saxena and Dhawan [43] produced more than 100 000 plants. Saxenaand Dhawan [43] noticed a strong genotypic effect at all stages of regeneration, thatis, initiation, multiplication, and germination of somatic embryos.Saxena and Dhawan [43] also studied various factors such as potting mixture,

humidity regime, and seasons, which are known to influence the transplantationprocess. The seasonal effect was found to be very pronounced. Saxena and Dhawan[43] encountered higher rates of mortality during winter (October–February). Theyattributed this to D. strictus being more susceptible to low temperatures because thespecies is well adapted to growth in warm areas. The tissue-cultured plants usuallytake 6–8 weeks after transplantation to form rhizomes. According to Saxena andDhawan [43], plants transplanted in July survived winter as the plants producedrhizomes, and plantlets transferred to soil in October or later died because theywere exposed to low temperatures before they formed rhizomes.Singh et al. [67] successfully regenerated NaCl-tolerant plantlets of D. strictus via

somatic embryogenesis from NaCl-tolerant embryogenic callus. Selection ofembryogenic callus tolerant to 100mM NaCl was made by exposing the callus


cultures to 0–200mM concentrations of NaCl in callus initiation medium (MSmedium supplemented with 13.6 mM 2,4-Dþ 2.3 mM Knþ 3% sucroseþ 0.8%agar). The NaCl-tolerant embryogenic callus differentiated somatic embryos onmaintenance medium (MSmediumþ 9.0mM2,4-Dþ 2.3mMKnþ 3% sucroseþ 0.8%agar) and containing 0–200mM NaCl. About 39% of mature somatic embryoswere tolerant to 100mM NaCl. These somatic embryos germinated and convertedinto plantlets in half-strength MS mediumþ 0.1 mM NAAþ 0.49 mM IBAþ 2%sucroseþ 100mM NaCl. About 31% of these plantlets established well on trans-plantation into a garden soil and sand (1: 1) mixture containing 0.2% (w/w) NaCl.These results suggested that plantlets regenerated from 100mM NaCl-tolerantembryos can survive on soils rich in salt.

13.6Future Prospects

D. strictus is a clump forming tropical bamboo adapted for growth in a wide rangeof climatic conditions. In D. strictus, methods are available for in vitro propagationthrough enhanced shoot multiplication from juvenile and mature explants, somaticembryogenesis, and in vitro induction of microrhizomes. In vitro selection proce-dure for developing salt-tolerant variant is also available. Developing a frost-tolerantvariant of this bamboo as well as developing drought-tolerant monopodial (lepto-morph) temperate bamboos, frost-tolerant sympodial (pachymorph) bamboos, andsalt tolerance in both the groups will be very useful in extending the range ofgrowth habitats of bamboos.

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Improving Crop Productivity in Sustainable Agriculture (TUTEJA:CROP IMPROVEMENT O-BK) || Bamboo: Application of Plant Tissue Culture Techniques for Genetic Improvement of Dendrocalamus