The Propagation of Species from Rooted Stem Cuttings Author(s): Ralph Lundquist and John G. Torrey Reviewed work(s): Source: Botanical Gazette, Vol. 145, No. 3 (Sep., 1984), pp. 378-384 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/2474733 . Accessed: 03/04/2012 15:54

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http://www.jstor.org BOT. GAZ. 145(3):378-384. 1984. ? 1984 by The Universityof Chicago. All rightsreserved. 0006-8071/84/4503-0005$02.00

THE PROPAGATION OF CASUARINA SPECIES FROM ROOTED STEM CUTTINGS

RALPH LUNDQUIST AND JOHN G. TORREY Cabot Foundation,Harvard University, Petersham, Massachusetts 01366

Juvenile,mature softwood, and maturehardwood stem cuttings from 1- to 6-yr-oldtrees of the Casuarina- ceae were testedfor their capacity to formroots. Trials includedmist-bench with bottom heat, water-culture jars, and, forlarge numbers,an aeratedhydroponic system. ,C. cunninghamiana, C. montana,C. suberosa, and Gymnostomapapuanum were all successfullyrooted. Greatest success (60%- 100%) was achievedwith mature softwood stem cuttings using auxin treatment (50 mg/literindole-3-butyric acid fora 3-h soak) followedby waterculture in l/4-strengthHoagland's solutionminus nitrogen. Inoculating 4-6-wk-old rootedcuttings with nodule suspensionsor pure culturesof HFPCcI3 gave 80%-89% nodule formationat 3 mo.

Introduction wood cuttingsof Casuarina equisetifoliaand C. Speciesof thegenus Casuarina have been widely junghuhnianausing growthsubstance treat- disseminatedby man and now occurthroughout the ment.With a view to producingclonal materialfor equatorialand subequatorialregions of the world, experimentalwork, we undertooka studyof the servingfor fuel wood, land reclamation,dune sta- conditionsfavoring rapid propagationof young Casuarina plantsfrom stem cuttings and theirnod- bilization,and shelterbelts (NATIONAL ACADEMY ulation. OF SCIENCES 1980, 1984; MIDGLEY et al. 1983). The plantsare wind pollinated;they and fruitthroughout their lives and have been propa- Material and methods gated largelyfrom . The roots of Casuarina SOURCE OF are readilyinfected by the soil bacteriumFrankia ()and producenodules that fix at- Three genera withinthe familyCasuarinaceae mosphericnitrogen (TORREY 1982). When were grownfrom seed. Plantswere establishedin are plantedin new sites, Frankia must be intro- greenhousesat the Harvard Forest, Petersham, duced in soil samples taken fromexisting nodu- Massachusetts,from various seed collections(table latedtrees or fromlitter containing old rootsor root 1). up to 6 yr old were grownsuccessfully nodules.Propagation by seed and modulationby soil in large containersin pottingsoil but requiredse- or leafylitter are provedmethods for establishing verepruning at regularintervals to keep theplants largeplantations of variousmembers of theCasua- withinthe greenhouseroof. Species of Casuarina rinaceae (MIDGLEY et al. 1983). were effectivelymodulated in the greenhousewith For selectionand geneticimprovement of spe- nodulesuspensions from field-collected nodules of cies of ,it is preferableto propagate C. equisetifoliacollected in Florida. Plants were selectedtrees vegetatively from superior specimens wateredregularly with tap water and at intervals and to achieve modulationwith strains of Frankia with l/4-strengthHoagland' s solution(HOAGLAND knownto be highlyinfective and to producenod- and ARNON 1950) minus nitrogen.Species of Al- ules withhigh rates of N2 fixation,thereby maxi- locasuarina and Gymnostomausually failed to mizingthe benefitsof the symbioticassociation. nodulatein our greenhousesand were sustained DIEM and DOMMERGUES (1983), DIEM et al. by regularfertilization with l/4-strengthcomplete (1983a, 1983b), and ZHANG et al. (1984) reported Hoagland's solution. thatpure culturesof Frankia isolated fromCasu- arina root nodules are available to use as inocu- TYPES OF STEM CUTTINGS lum. On theother hand, published accounts of veg- Three typesof cuttingswere testedfor rooting etativepropagation are sparse (cf. TORREY 1983), capacity. "Mature cuttings"taken fromtrees be- and the older methodsreported, including air lay- tween1 and 6 yrold wereeither "softwood" (green eringand rootingof cuttings,are relativelycrude stem)or "hardwood"(brown stem and woody) cut- and unpredictable,with low yield. SOMASUN- tings.In some species, 3-4-yr-oldtrees were cut DARAM and JAGADEES (1977) and HUSAIN and PON- offat the main trunkca. 10-20 cm above the soil NUSWAMY (1980) reportedsuccess in rootingsoft- level. These trunksregenerated many new shoots Manuscriptreceived October 1983; revised manuscriptre- fromthe stump. Cuttings taken from these regrown ceived February1984. shoots were termed"juvenile cuttings."Experi- Addressfor correspondenceand reprints:JOHN G. TORREY, mentson rootingcuttings were performed through- Cabot Foundation,Harvard University, Petersham, Massachu- out all seasons overa periodof severalyears, using setts01366. ambientgreenhouse conditions.

378 LUNDQUIST & TORREY-PROPAGATION OF CASUARINA 379

TABLE 1

SCIENTIFIC NAMES AND SOURCES OF SEEDS USED FOR GREENHOUSE PLANTINGS

Plant species Source Donor Division of Forest Research, CSIRO, Casuarina cunninghamiana ...... Canberra, D. BOLAND Clear Lake, Florida; W. SILVER C. equisetifolia...... Hilo, Hawaii R. SKOLMEN C. montana(= C. cunninghamiana) ...... Maui, Hawaii P. WOOMER Lyon Arboretum, C. suberosa (= Allocasuarina littoralis) ...... Honolulu, Hawaii R. BAKER King's Park Botanical Garden, Allocasuarina lehmanniana ...... Perth,W. Australia DIRECTOR OF Waiakea Arboretum, THE PARK Gymnostomapapuanum ...... Hilo, Hawaii R. SKOLMEN

NoTE.-Nomenclature follows JOHNSON (1982).

METHODS OF PLANT GROWTH SUBSTANCE to theholes in theperforated plastic, and theirbases TREATMENT wereimmersed in a commonsolution of l/4-strength Hoagland's solution at In earlyexperiments, cuttings ca. 7 cm long were minusnitrogen pH 7. excised fromshoot tips and dipped into commer- INOCULATION OF ROOTED CUTTINGS cial rootingcompounds including Rootone or Hor- modin,which contain different levels of indole-3- Once successfullyrooted, cuttingswere trans- butyricacid (IBA). These cuttingswere thenf placed ferredto potsor othercontainers for establishment. in sand in flats,in a rootingbed, or in watercul- Upon transfer,the rootedcuttings were inoculated ture. In later more extensivetrials, several plant with microbialpreparations of the actinomycete growthsubstances were testedat a range of con- Frankia,which induces N2-fixing nodules on roots centrations.Synthetic substances tested included: by root hair infection(CALLAHAM et al. 1979). indole-3-aceticacid (IAA), IBA, and c.-naphtha- Suspensionswere preparedby grindingin distilled leneacetic acid (NAA). Most of these trials in- waterwith a mortarand pestlefresh whole nodules volveda 3-hdip in auxinsolution; in others,shoots fromgreenhouse-grown plants. Roots were im- were placed in nutrientsolution with added auxin mersedin thesuspension before transfer of thecut- forlong-term treatment. In bothcases rootingwas tingsto sand in pots. Later, whenpure culturesof performedin l/4-strengthHoagland's solutionmi- Frankiaisolates were available (ZHANG et al. 1984), nus nitrogenat pH 7. bacterialcultures were centrifuged,washed in in- organicsolution to removeany combined nitrogen, HANDLING THE TREATED CUTTINGS homogenized,and thenapplied directly to theroots or to the stembase of establishedcuttings in sand In theearly experiments, auxin-treated stem cut- in pots. Successfulmodulation following inocula- tingswere plantedin sand beds in the greenhouse tionwas determinedby directexamination for root andwere provided bottom heat and intermittentmist. noduleson theroot systems after several weeks or A secondprocedure involved rooting cuttings in by observing the successful establishmentand water-culturejars: 8-oz softglass jars wrappedin growthof theinoculated cuttings when watered with foilwere capped withlids coveredwith perforated nutrientsolution lacking combined nitrogen. opaque plasticfilm. The basal halves of 7-8-cm- long cuttingswere treatedwith auxin solutionand Results then transferredto jars containing l/4-strength Hoagland's solutionwithout nitrogen at pH 7 (fig. EXPERIMENTS WITH MATURE HARDWOOD CUTTINGS 1). The basal ends were examinedat intervalsand Since we did nothave access to fullymature trees responses recorded. Rooted cuttingswere later growingin thefield, experiments on therooting of transferredto pots (fig. 2). stemcuttings from fully grown trees were limited. A thirdtechnique involved trough culture. With In general,our experiencewas thatolder stem cut- a view to handlinglarger numbers of cuttingswith tingswere more difficultand took longerto root singletreatments, we experimentedwith a simple, thanjuvenile materialbut thatboth maturesoft- aerated, polyethylene-coveredplastic trough in wood and hardwoodcuttings could be rootedwith whichseveral hundred cuttings could be rootedsi- appropriateauxin treatment.Response to auxin multaneously(figs. 3, 4). Afterauxin treatment and treatmentwas slower in maturecuttings, and the a tap waterrinse, cuttings were transferredsingly species influencedthe responsemarkedly. ~~~~,~~~~~~II~~~~~~~~~I~~~~~~~ui~~~~~~~~~L ______~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~IIIIII

FIGS. 1-4. -Fig. 1, Mature softwoodstem cuttings of Casuarinaequiseti~folia placed in waterculture for rooting experiments on the effectsof auxin treatment.Containers were 8-oz soft glass jars covered with foil. Lids were perforated,opaque plastic sheets held in place with cardboardrings. Fig. 2, Rooted cuttingsof Casuarinaspecies pottedafter inoculation and grown in sand-vermiculitein 3-inch pots. Fig. 3, Stem cuttingsof C. cunninghamianarooted in shallow troughwith aeration of nutrient solution. Fig. 4, Invertedtop of troughin fig. 3 withrooted cuttings in place, showingextensive rooting achieved in the aerated solution. Small black tube is the aerationline. LUNDQUIST & TORREY-PROPAGATION OF CASUARINA 381

TABLE 2

PRELIMINARY EXPERIMENTS ON ROOTING AND CALLUS FORMATION OF MATURE HARDWOOD STEM CUTITINGS IN A SAND BED WITH PERIODIC MISTING IN THE GREENHOUSE

Species, no. of cuttings, Callused Rooted and durationof experiment Treatment' (%) (%)

Casuarina equisetifolia: 20, 5 wk ...... Rootone-F 60 0 20, 5 wk ...... Untreated 20 0 20, 8 wk ...... Rootone-F 10 90 15, 6 wk ...... Untreated 40 27 15, 6 wk ...... 50 mg/literNAA 20 47 15, 6 wk ...... 50 mg/literIBA 60 20 15, 6 wk ...... 50 mg/literIAA + 50 mg/literIBA 0 0 C. montana. 25, 4 wk ...... 50 mg/literIBA 4 0 25, 4 wk ...... 50 mg/literNAA 0 0 25, 4 wk ...... 50 mg/literIBA + 50 mg/literNAA 0 0

aRootone-F was used as a talc dip; auxins were given as a 3-h soak beforeplacing cuttingsin sand bed.

Resultsof some preliminarytrials with mature mist-benchconditions, the plantswere more sus- hardwoodstem cuttings,some treatedwith com- ceptibleto fungalattack, and mistingwas regarded mercialroot compounds in talc as a dip and others as bothunnecessary and unfavorableand was dis- withpure plant growth substance treatments using continued. a 3-h soak, were variable (table 2). Casuarina equisetifoliaappeared to be most responsive to ROOTING IN WATER CULTURE treatment.Callus formationusually accompanied Among the Casuarina species tested in water any successfulrooting response. Because of early culture,the mature softwood cuttings failed to root success in rootingjuvenile and maturesoftwood withoutauxin treatment,although some cuttings stemcuttings, work on older hardwoodstem cut- formedcallus (table 4). IBA seemed to be consis- tingswas discontinued. tentlybetter than NAA; 1AA was nottested in these trialsas it was consistentlyless effective(table 3). JUVENILE AND MATURE SOFTWOOD MATERIAL Amongthe various species testedunder a wide rangeof conditionsand treatments,softwood stem TABLE 3 cuttingswere most easily rooted and establishedas ROOTING AND CALLUS FORMATION ON STEM CUTTINGS OF new plants.Our trialsto determinethe most effec- CASUARINA SPECIES PROPAGATED IN SAND BED tive root-inducingauxin, its concentration,and WITH PERIODIC MISTING IN THE GREENHOUSE methodof applicationwere most extensiveon ju- venileand maturesoftwood material. C. EQUISETIFOLIA C. MONTANA Softwoodcuttings show two typesof regenera- (mature softwood)a (juvenile)a tive response:callus formationand/or root initia- TREATMENT Callused Rooted Callused Rooted tion. In manytreatments, only callus was formed; (mg/liter) (%) (%) (%) (%)

in others,callus formationwas followedby root Water control . . 93 7 7 93 initiation,usually at otherthan callus sites.In other IAA: treatments,roots formed without callusing. 25 ...... 40 0 0 100 50 ...... 73 7 0 100 100 ...... 40 20 7 73 ROOTING IN SAND ON MIST-BENCH IBA: The juvenilecuttings from C. montana(possibly 25 ...... 53 20 7 93 C. cunninghamianaor withclose affinityto it) were 50 ...... 40 47 0 93 100 ...... 53 27 7 40 easily rootedand showed littlebenefit from auxin NAA: treatment(table 3). Callusingwas infrequent.Ma- 25 ...... 33 20 0 100 ture softwood cuttingsof C. equisetifolia re- 50 ...... 27 60 0 100 spondedwell to appropriateconcentrations of auxin 100 ...... 13 0 20 100

withoptimum results with IBA or NAA at 50 mg/ a Mature softwood or juvenile stem cuttings ca. 7 cm long liter(table 3). Callusingwas abundanteven with- were placed with basal 2 cm in treatment solutions for 3 h out auxin treatmentand tendedto be repressedby before planting in the sand bed, 15 cuttings/treatment. Data concentrationsfavorable to root initiation.Under collected after 35 days. 382 BOTANICAL GAZETTE [SEPTEMBER

TABLE 4

ROOTING AND CALLUS FORMATION ON STEM CUTTINGS OF SPECIES OF CASUARINACEAE PROPAGATED IN WATER CULTURE JARS IN THE GREENHOUSE

CASUARINA C. EQUISETIFOLIA' CUNNINGHAMIANA' GYMNOSTOMA PAPUANUMa C. SUBEROSA (putative)a Callused Rooted Callused Rooted Callused Rooted Callused Rooted TREATMENT (%) (%) (%) (%) (%) (%) (%) (%) (mg/liter) (25 days) (42 days) (18 days) (18 days) (30 days) (14 days) (14 days) (30 days)

Watercontrol ... 93 0 0 0 73 0 47 0 0 0 IBA: 25 ...... 20 60 7 3 7 67 80 0 30 60 50 ...... 20 60 0 27 0 73 80 10 40 80 100 ...... 7 20 40 13 13 73 80 0 40 90 NAA: 25 ...... 7 27 0 27 7 53 80 9 36 55 50 ...... 33 33 13 0 13 60 60 10 60 70 100 ...... 7 27 20 7 33 47 67 0 30 50

a Mature softwoodstem cuttingsca. 7 cm long were given 3-h soak in auxin solution,transferred to l/4-strengthHoagland's solutionminus nitrogen at pH 7; 15 cuttings/treatment.Measurements recorded after the numberof days shownfor each species.

Casuarina suberosa was inducedto rootat a range INOCULATION TRIALS WITH ROOTED CUTTINGS of concentrationsof bothIBA and NAA withmax- Rootedcuttings of Casuarina usuallyformed one imum effectivenessat 50 mg/liter.Gymnostoma or two majorvigorous adventitious roots, which in papuanum cuttingsrooted withouttreatment, but turnformed numerous short lateral roots. When theresponse could be increasedand accelerated with placed in pots, the mainroots were bentaround to plantgrowth substance treatment (figs. 5, 6). This fitthe pot (fig. 2). Successful inoculationof the species formedboth callus and roots (figs. 7, 8) rootscould be achieved by dippingthe roots into withoutauxin treatment. a nodule suspensionor a suspensionof pure cul- ROOTING OF CUTTINGS IN TROUGHS turedFrankia. Using two separateseries of rooted cuttingsfrom two differentplants of C. equiseti- Large numbersof cuttings,as would be needed folia, inoculationwith nodule suspensions and with forclonal propagationof a specimentree, could be Zhang'sisolate HFPCcI3 resultedin modulationrates rootedin hydroponictroughs (figs. 3, 4). As in of 83% and 89%, witha mean numberof nodules earlierexperiments with C. equisetifolia,basal cal- of 4.1 in the firstseries and 10.7 in the second. lusingoccurred spontaneously without auxin treat- Numbersof nodules per plant at 3 mo afterinoc- ment(table 5). Rootingresponses were best at low ulationvaried from one perplant, usually quite large, auxinconcentration. Callus formationtended to be to 37 nodules per plant (each quite small). Acet- suppressedby auxin treatmentin this species (see ylenereduction trials indicated these nodules were also tables 3, 4). effectivein fixingdinitrogen, and the survivaland growthof themodulated cuttings confirmed this re- TABLE 5 sult. Rooted and modulatedcuttings could be es- ROOTING AND CALLUS FORMATION ON STEM CUTTINGS OF tablishedreadily and in good numbersby these CASUARINA EQUISETIFOLIA PROPAGATED IN TROUGH methods. CULTURE IN THE GREENHOUSE

Treatment (mg/liter) Callused (%)a Rooted (%) Discussion

Water control ...... 80 7 These experimentslead to the recommendation IBA: of a simple,inexpensive, and effectivemethod for 25 ...... 7 80 the vegetativepropagation of softwoodstem cut- 50 ...... 13 60 tingsof species of the Casuarinaceae. Young soft- 100 ...... 0 40 NAA: wood cuttingstaken from the growing parts of ma- 25 ...... 0 53 ture trees of Casuarina spp. respond to auxin 50 ...... 0 53 treatmentwith callus and/orroot initiation.IBA, 100 ...... 7 40 the common componentof commercial rooting powders,was themost effective of theplant growth a Mature softwood stem cuttings ca. 7 cm long were given a 3-h soak in auxin solution, rinsed, and placed in trough with substancestested. The concentrationand the du- '/4-strengthHoagland's solution minus nitrogen at pH 7 with rationof treatmentfor optimum response should be aeration; 15 cuttings/treatment.Measured after 30 days. determinedfirst in small lots of cuttings,prefera- 19841 LUNDQUIST & TORREY-PROPAGATION OF CASUARINA 383

..' ...... s sno -| ~~~~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...... | Iw x~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...... HE ,.w

' "1" 1 w j ...... ,,,!II!!!llsl.!,!,..!_-is...... ,. ...., d o~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...... ^;Y 1~~...... AA 5PP

Bl'v! ...... # ~ ~ ~~~~ ~ ...... ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...... I6 (D~~~~~~~~~~~~~~~~. ..,N 1 .

..../......

FIGS.~~~~~~~~...... 5,...... Roote aua .... ftrnary6 ...... ki ntintsltini wtrcutr jars.~~~~~~~~~~~~~~.mg/literSeeligscaluedan.rotd.itou..xiNA before ransferp.p.....in.fig towater-culure jars...... Fg.,Rotdse.utig.sinfgig. 7, Clser view o callus ad roots ofcuttings utwtf G. hsa n2 6. Fig. 8, Details of callus formation on stem cuttings of G. papuanum nearly 6 wk after treatment.formation~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.::Swail Note that callus is~~~~~~~~..tic ..o~~~~ formation......

blyin water-culture jars; then large samples can be tothree robust elongating roots with many laterals. handledefficiently and effectively.No specialar- These rootscan be inoculatedeffectively with rangementsfor bottom heating or intermittent mist- Frankia,using suspensions either from pure cul- ingare required. turesor fromground nodule preparations by dip- Rootedcuttings after 4-6 wktypically bear two ping.The cuttingscan be transplantedto pots or, 384 BOTANICAL GAZETTE in large numbers,to conical tubes or Rootrainers effectivelyseedlings of C. cunninghamiana,C. wherethe modulatedseedlings develop rapidly. equisetifolia,C. glauca, and C. junghuhniana.The Earlierstudies reviewed by TORREY(1983) were same methodscould be applied to rootedcuttings. ratherpessimistic about methodsfor propagating Thus, thereare now available suitablemethods Casuarina. Air layering,whether of large or small forpropagation from stem cuttings of a numberof branches,was slow and inefficient,producing low Casuarina spp., using plant growth substance yields (SYED 1957; CHAUDHARY and RAM 1961). treatmentand inoculatingrooted cuttings with se- Using long "tender"shoots, SOMASUNDARAM and lected pure culturesof the effectiveactinomycete JAGADEES (1977) found plant growth substance Frankia. Fromour studies(ZHANG et al. 1984) and treatmentessential for successfulrooting of stem fromthose of DIEM et al. (1983a), othergenera of cuttings,and HUSAIN and PONNUSWAMY(1980) used the Casuarinaceae, i.e., Allocasuarina and Gym- shootand root suckers for propagation of male plants nostoma(cf. JOHNSON 1982), mayrequire still dif- of C. junghuhniana,for which seeds were un- ferentstrains of Frankia thanhave proved useful available. with Casuarina. Until these are isolated, nodule KONDAS(1983) describedmethods of vegetative suspensionsfrom the appropriatehost plant should propagationof C. junghuhnianafor mass plantings proveeffective in inducingroot modulation. in , using 10-cm-longstem pieces in red loam undermist tents.In 45 days rootingsuccess fol- lowingtreatment with rooting substances varied from Acknowledgments 40% to 90%, dependingon thetime of year. CHIT- This researchwas supportedin partby theMaria TACHUMNONK(1983) reportedexperiments in Thai- Moors Cabot Foundationfor Botanical Research of land on theeffectiveness of IBA treatmentin root- HarvardUniversity. J. G. TORREY expresses his ingstem cuttings of C. junghuhniana.Cuttings were appreciationto thedonors listed in table 1 fortheir given 24-h auxin treatment,planted in polyeth- generousassistance in makingfield collectionsof ylene bags containingtopsoil, and maintainedat seeds. We acknowledge the assistance in the high humidityfor 30 days beforehardening and greenhouseof SHIRLEY LAPOINTE and expressap- planting. preciationto FRANCES O'BRIEN and DOROTHY SMITH DIEM et al. (1983a) reportedprocedures for in- forsecretarial assistance. Thanks are expressedto oculating seeds of C. junghuhniana applied in ZHANG ZHONGZE for his cooperationwith the in- nurseryplantations in Senegal. Their isolate of oculationtrials. We are also indebtedto W. OR- Frankia strainORS 1106 was found to nodulate MEROD and S. LANCELLE forphotography.

LITERATURE CITED

CALLAHAM,D., W. NEWCOMB,J. G. TORREY,and R. L. PE- crop in India. Pages 66-76 in S. J. MIDGLEY,J. W. TURN- TERSON.1979. Root hair infectionin actinomycete-induced BULL, and R. D. JOHNSTON, eds. Casuarina ecology, man- rootnodule initiation in Casuarina, Myrica, and Comptonia. agementand utilization.CSIRO, Melbourne. BOT. GAZ. 140(suppl.):S1-S9. MIDGLEY, S. J., J. W. TURNBULL, and R. D. JOHNSTON, eds. CHAUDHARY,N. R., and T. RAM. 1961. Common methodsof 1983. Casuarina ecology, management and utilization. vegetativepropagation as usefulin forestry.Proc. TenthSil- CSIRO, Melbourne.286 pp. vicult.Conf., pp. 214-223. Dehra Dun, India. NATIONALACADEMY OF SCIENCES.1980. Firewood crops. Na- CHI-rACHUMNONK,P. 1983. Silvicultureof Casuarinajung- tionalAcademy Press, Washington,D.C. 237 pp. huhnianain . Casuarina ecology, managementand . 1984. :nitrogen-fixing trees for adverse sites. utilization.CSIRO, Melbourne. National Academy Press, Washington,D.C. 118 pp. DIEM, H. G., and Y. DOMMERGUES.1983. The isolation of SOMASUNDARAM,T. R., and S. S. JAGADEES.1977. Propagation Frankia fromnodules of Casuarina. Can. J. Bot. 61:2822- of Casuarina equisetifoliaForst. by plantingshoots. Indian 2825. Forest. 103:737-738. DtEM,H. G., D. GAUTHIER,and Y. R. DOMMERGUES.1983a. SYED, H. 1956. A note on vegetativemultiplication of Casu- Inoculationof Casuarina using a pure cultureof Frankia. arina cunninghamiana.Indian Forest. 83:226-227. NitrogenFixing Res. Rep. 1: 18-19. TORREY, J. G. 1982. Casuarina: actinorhizalnitrogen-fixing 1983b. An effectivestrain of Frankia fromCasuarina tree of the tropics. Pages 427-439 in P. H. GRAHAM and sp. Can. J. Bot. 61:2815-2821. S. C. HARRIS,eds. Biological nitrogenfixation technology HOAGLAND,D. R., and D. I. ARNON. 1950. The water-culture fortropical agriculture. CIAT, Cali, Colombia. methodfor growing plants without soil. CaliforniaAgr. Exp. . 1983. Root developmentand rootmodulation in Casu- Sta. Circ. 347. arina. Pages 180-192 in S. J. MIDGLEY, J. W. TURNBULL, HUSAIN,A. M. M., and P. K. PONNUSWAMY.1980. Propagation and R. D. JOHNSTON, eds. Casuarina ecology, management of by planting shoots and root and utilization.CSIRO, Melbourne. suckers.Indian Forest. 106:298-299. ZHANG ZHONGZE, M. F. LOPEZ, and J. G. TORREY. 1984. A JOHNSON,L. A. S. 1982. Notes of Casuarinaceae. II. J. Ade- comparison of cultural characteristicsand infectivityof laide Bot. Gardens6:73-87. Frankiaisolates from root nodules of Casuarina species.Plant KONDAS,S. 1983. Casuarina equisetifolia-a multipurposecash and Soil 78:79-90.