Evaluation of uva camarona (Macleania rupestris Kunth A.C. Smith) propagation with air layering

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1 PROPAGATION AND TISSUE CULTURE Evlution of uv cmron (Mcleni rupestris Kunth A.C. Smith) propgtion with ir lyering Evlución de l propgción de uv cmron (Mcleni rupestris Kunth A.C. Smith) por medio de codos éreos Sntigo Durán-Css 1, Clr Veloz-Sun 1, Stnislv Mgnitskiy 1, nd Héctor Orlndo Lncheros 2 ABSTRACT Uv cmron (Mcleni rupestris Kunth A.C. Smith) elongs to the Ericcee fmily nd grows in pármo nd supármo res in Colomi, etween 2,200 nd 3,500 m.s.l. This plnt presents edile erries tht serve s source of food nd smll income for locl communities. The sence of propgtion protocol for this species limits its use. This study imed to evlute the effectiveness of sexul propgtion of M. rupestris with ir lyering, using indole-utyric cid (IBA) s rooting hormone t different concentrtions: 0, 500, 1,000, nd 2,000 mg L -1. The results showed tht n exogenous ppliction of IBA ccelerted the rooting process in the lyered zone, with notle emission of first dventitious roots t 60 dys from the strt of the experiment. The tretments of 500 nd 1,000 mg L -1 IBA hd the highest numer of roots per lyer, eing two to three times higher thn those presented in the control. No significnt differences were seen in root length etween tretments. The tretments of 500 nd 1,000 mg L -1 IBA showed high production for the dry weights of the roots nd cllus, with higher weight of cllus compred to root weight. Air lyering negtively ffected the longitudinl growth of the rnches, since their verge growth rte ws 1.49 cm per month, while the growth of intct rnches ws 2.78 cm per month. The results suggest tht the est concentrtion for rooting ws mg L -1 IBA ecuse it hd the lrgest numer nd dry weight of roots in ir-lyered M. rupestris. Key words: Ericcee, indole-utyric cid, vegettive propgtion, roots, cllus. RESUMEN L uv cmron (Mcleni rupestris Kunth A.C. Smith) es un plnt de l fmili Ericcee que crece en zons de pármo y supármo en Colomi, entre los y msnm. Present ys comestiles, ls cules sirven como limento o fuente de pequeños ingresos pr ls comuniddes locles. L usenci de un protocolo de propgción de est especie limit su provechmiento. El presente trjo tuvo el ojetivo de evlur l eficci de l propgción sexul medinte codos éreos, utilizndo como hormon de enrizmiento ácido indolutírico (AIB) diferentes concentrciones: 0, 500, y mg L -1. Los resultdos mostrron que l plicción exógen de AIB celeró el proceso de enrizmiento en l zon de codmiento, oservndo emisión de ls primers ríces dventicis los 60 dís de inicido el ensyo. Los trtmientos de 500 y mg L -1 de AIB presentron el myor número de ríces por codo, siendo dos tres veces myor que ls presentes en el control. No huo diferencis significtivs en cunto longitud de ríces entre los trtmientos. L myor producción de peso seco de ríces y de cllo se presentó en los trtmientos de 500 y mg L -1 de AIB, mostrndo un myor peso de cllo comprdo con el de ríces. El codmiento fectó negtivmente el crecimiento longitudinl de ls rms, y que l ts de crecimiento promedio de ésts fue de 1,49 cm por mes, mientrs que ls que no se codron presentron un crecimiento de 2,78 cm por mes. Los resultdos sugieren que con l concentrción proximd de mg L -1 de AIB se otienen el myor número y peso seco de ríces en los codos de M. rupestris. Plrs clve: Ericcee, ácido indolutírico, propgción vegettiv, ríces, cllo. Introduction Within the gret diversity of plnt species of high use potentil in Colomi, Anden fruit trees re found, with only few domesticted species nd the mjority grown in the wild. These plnts ccount for n importnt source of food nd/or income for locl communities (Ntionl Reserch Council, 1989; Lgos-Burno et l., 2010). Mny of these fruit species re gining vlue, given their high commercil potentil due to the excellent fruit qulity for direct consumption (Vásquez nd Coimr, 2002; Received for puliction: 28 August, Accepted for puliction: 29 Mrch, Deprtment of Agronomy, Fculty of Agronomy, Universidd Ncionl de Colomi. Bogot (Colomi). sdurnc@unl.edu.co 2 Reserch Division, Jrdín Botnico José Celestino Mutis. Bogot (Colomi). Agronomí Colomin 31(1), 18-26, 2013

2 Snjinés et l., 2006) nd incresing demnd in North Americ, Europe, nd Jpn, where they re known s exotic Anden fruits (Lgos-Burno et l., 2010). The Ericcee fmily is one of the most representtive in the Anden forests, minly in mountin cloud forests, s well s pármo nd supármo regions, where it plys n importnt ecologicl role (Luteyn nd Pedrz-Peñlos, 2008; Lgos-Burno et l., 2010) s wter regultor, food source nd, in some cses, source of income for the inhitnts of the region (Lgos-Burno et l., 2010). Mcleni rupestris Kunth A.C. Smith is one of the undomesticted fruit species of the Ericcee fmily. This plnt is found in wild res of the Anden region t ltitudes etween 1,500 nd 4,100 m (Acero nd Bernl, 2003; Luteyn nd Vidl, in press). In Colomi, M. rupestris is commonly clled uv cmron or cmroner nd is locted in wild mountin hitts s constituent of ntive vegettion (Sturm nd Rngel, 1985; Lgos-Burno et l., 2010). This species is found in the provinces of Antioqui, Boycá, Clds, Cundinmrc, Huil, Met, Nriño, Norte de Sntnder, Putumyo, Risrld, Sntnder, nd Mgdlen in the Sierr Nevd de Snt Mrt, growing etween 2,200 nd 3,500 m (Crdozo et l., 2009; Lgos- Burno et l., 2010). Mcleni rupestris is n epithytic shru mesuring from 0.6 to 2.5 m tll with rnching originting lmost t the se of the stems (Acero nd Bernl, 2003; Crdozo et l., 2009; Lgos-Burno et l., 2010). Sometimes, this plnt grows ner tll trees nd scends through their rnches imitting vine species. The stems of M. rupestris re wvy nd hve cler nd scly rk. The young leves re red nd light green when mture (Fig. 1A), nd re lternte, simple, shiny, nd fleshy (Crdozo et l., 2009). The plnt hs xillry inflorescences locted t the pex of the rnches, without picl flowers; the inflorescences re rcemose with four to 20 flowers. The flowers hve ell-shped clyx lim typicl of the Ericcee fmily; the perinth of the flower nd its pedicel re pink to red (Fig. 1B), with cylindricl, succulent sepls (De Vlenci nd De Crrillo, 1991; Crdozo et l., 2009; Lgos-Burno et l., 2010). This species grows in rocky res, forests, nd ushes s prt of ntive vegettion. It is resistnt to strong winds nd frost, nd requires good sun exposure (Rodríguez nd Peñ, 1984). The fruit is edile nd origintes from n inferior ovry (Font Quer, 1965); it is stlked, nd erry type. Its color during the ripening process chnges from light pink, ple green, nd whitish to drk purple nd lmost lck in senescence (Fig. 1: C nd D), so the skin of the fruit is purple in mturity nd the pulp hs light color similr to grpe. The fruit is fleshy, hving dimeter of cm nd n verge weight of 2.6 g (De Vlenci nd De Crrillo, 1991; Acero nd Bernl, 2003; Crdozo et l., 2009). The fruits re rrnged on rcemose xis with cropetl mturtion similr to tht oserved in the flowers. The flowers re pendulous, ut the fruits re not, which indictes tht the twisting of petioles occurs during fruit development (De Vlenci nd De Crrillo, 1991). Bromtologicl chrcteriztion of M. rupestris fruits y Amy nd Rincon (1989) showed tht they hve totl cidity of 0.49% citric cid, 12.5 Brix solule solids, 1.8% crude fier, 1.3% crude protein, nd 0.08 g tnnin in 100 g of pulp. The contents of minerl elements per 100 g of pulp were mg clcium, 0.97 mg iron, 5.24 mg potssium, 8.69 mg sodium, nd 10.3 mg phosphorus; nd the sugrs (per 100 g of pulp): 15.9 g fructose, 1.62 g of sucrose, 13.5 g of α-glucose, nd 16.1 of β-glucose (Amy nd Rincon, 1989). Mcleni rupestris plnts hve on verge of 14 fruits per inflorescence, with two peks of production per yer in Colomi (De Vlenci nd De Crrillo, 1991), the first one from mid-jnury to April, nd the second one from August to mid-novemer. These periods fluctute with rinfll distriution; nd with dditionl rinfll in dry wether, the periods of flowering nd fruiting overlp (Crdozo et l., 2009). Humn ctivities hve led to out 75% of forestlnd locted in the northern Andes eing trnsformed to the point of totl deforesttion (Henderson et l., 1991). These ecosystems re home to out 80% of the different neotropicl Ericcee species nd some of them could e source of income for locl communities. For this reson, reforesttion progrms of these environments re needed, employing these species, tking into ccount tht mny Ericcee plnts re successful colonizers of nturl or nthropogeniclly distured res (Luteyn, 2002). In this sense, the potentil use of M. rupestris is not only limited to its use s commercil crop, ut lso to lnd restortion. The sence of n dequte M. rupestris propgtion protocol, tht is ffordle nd ccessile to smll growers is one of the ostcles to the introduction of this plnt into commercil cultivtion nd its use for the recovery of pármo nd supármo ecosystems. Therefore, it is necessry to undertke reserch tht seeks to procure consistent nd Durán-Css, Veloz-Sun, Mgnitskiy, nd Lncheros: Evlution of uv cmron (Mcleni rupestris Kunth A.C. Smith) propgtion... 19

3 relile plnt mteril of M. rupestris through the evlution of vrious methods of plnt propgtion. Air lyering is n ncient method used for vegettive propgtion of woody plnts tht cn ensure the sexul propgtion of species when sexul reproduction is very slow or ineffective (Adrince nd Brison, 1955), s is the cse of M. rupestris (Acero nd Bernl, 2003; Crdozo et l., 2009), or when there is no detiled protocol for propgtion through cuttings (Adrince nd Brison, 1955). The methodology of ir lyers cn otin 100% rooting ecuse the rnch is rooted to the mother plnt; in ddition, homogeneous plnts re otined within reltively short period of time (Adrince nd Brison, 1955; Lme et l., 1990). In the literture, no dt re found on the possile propgtion of M. rupestris with ir lyering, which would e new method for this species. Therefore, this study imed to evlute the possiility of sexul propgtion of M. rupestris y ir lyering, in order to provide the tools nd methodologies tht would fcilitte the commercil propgtion nd use of this species. Mterils nd methods The study ws crried out in the period from August to Decemer 2011, in the collection of pármo plnts of the José Celestino Mutis Botnicl Grden, Bogotá (Colomi), locted t the coordintes N nd W, nd ltitude 2,552 m.s.l. During this period, the following were registered: mm of cumultive rinfll, verge temperture of 18 C, verge photosyntheticlly ctive rdition of μmol m -2 s -1 nd wind speed of 0.14 km h -1 (Sutron wether sttion, José Celestino Mutis Botnicl Grden). The vegettion in ssocition with Mcleni AC rupestris Kunth Smith plnts ws typicl of the pármo nd supármo ecosystems of Colomi, with species elonging to the fmilies: Ericcee, Melstomtcee, Astercee, mong others. The sndy lom soil where the M. rupestris plnts were estlished hd n verge ph of Adult individuls of M. rupestris were selected with ctive vegettive growth, locted under similr full light exposure, nd without visul symptoms of nutritionl deficiencies or diseses. The selected plnts hd pproximtely 10 to 15 stems nd n verge height of 2.5 m. The lyering ws performed on six individul plnts, mking four ir lyers per plnt, with ech lyer corresponding to specific tretment; the rnches selected hd one lyer per rnch. Therefore, in totl, six replictes were estlished for ech tretment, so tht every plnt hd full set of tretments. Lignified orthotropic stems t the vegettive growth stge (with presence of few flowers, if ny, nd without the presence of fruits) were chosen on every selected plnt, with homogeneous chrcteristics, presenting turgid leves nd vigorous growth. Selected rnches underwent complete ring cut in the middle of the rnch, removing 4.0 cm portion of the rk with grfting knife previously disinfected with 0.5% sodium hypochlorite. The ppliction of the rooting hormone ws performed immeditely fter removing the rk. With the id of rush, the ring ws homogeneously covered with the powder mixture, which contined indoleutyric cid (IBA) (99% purity, Biopck - Buenos Aires, Argentin) nd technicl tlc s n inert mteril in proportions tht yielded four tretments: 0, 500, 1,000, nd 2,000 mg L -1 IBA. After ppliction of the hormone, 200 g of previously moistened pet moss without nutrients (Klsmnn-Deilmnn GmH, Geeste, Germny) were rrnged round the cut tht ws further covered with trnsprent nd lck plstic, thus, voiding light nd moisture loss. Biweekly irrigtion ws performed with 5 ml of deionized wter on ech lyering, using syringe. Monthly, the length of the rnches ws mesured with mesuring tpe (from the top of the girdling zone to the pex of the rnch) nd the dimeter of ech rnch ws recorded with lim cliper 30 cm from the top of the lyer (ccurcy ±0.01 cm). The lyered rnches hd n verge length of 67 cm nd width of 1.1 cm when the experiment ws estlished. Furthermore, monitoring of root emission on ech lyer ws performed fter 60 d from the strt of the experiment. The mesurement of quntittive vriles ws performed t the Lortory of Plnt Physiology, Fculty of Agronomy of the Universidd Ncionl de Colomi, Bogot. This took plce 120 d fter the strt of the experiment, performing destructive smpling of ll lyered rnches. The cllus nd dventitious roots were removed using sclpel, weighed immeditely using high precision nlyticl lnce (±0.001 g, Ohus, Ohus Corp., Pine Brook, NJ) nd the dry weight ws determined fter plcing the smples in oven t 65 C for 48 h. The vriles mesured were: numer nd length of dventitious roots, fresh nd dry weights of roots nd cllus. A photogrphic record of the results ws lso performed. The sttysticl nlysis of the results ws crried out with the help of SAS version 9.0 nd the differences mong the 20 Agron. Colom. 31(1) 2013

4 50 40 y=-2e -05 x x R 2 =0.995 B Roots numer C IBA concentrtion (mg L -1 ) A FIGURE 1. Mcleni rupestris plnt.. Leves.. Inflorescence. c. Immture fruit. d. Mture fruit. Photos: Veloz-Sun, C. nd S. Durán-Css. E FIGURE 2. Mcleni rupestris root numer in ir lyerings t different IBA concentrtions. Tretments with the sme letters re not significntly different ccording to Tukey test (P 0.05). The rs indicte stndrd devition. D 2,000 mg L -1 1,000 mg L mg L -1 0 mg L -1 1 cm 1 cm FIGURE 3. Rooting of M. rupestris rnches t different IBA concentrtions. Photos: Veloz- Sun, C. nd S. Durán-Css. tretments were evluted y ANOVA with significnce level of 95% (P 0.05). The polynomil regressions were dditionlly determined mong the vriles. Results nd discussion At 120 d fter the strt of the experiment, 100% rooting for ll tretments ws found, without ny ded lyered rnches. This confirms the dt descried y Hrtmnn et l. (2010), who indicted tht the lyering technique is successful in propgting plnts ecuse the lyered rnch is not seprted from the mother plnt nd, therefore, continues to receive wter nd nutrients. These dt gree with the results found in ir-lyers of grz (Vccinium meridionle Swrtz) tht presented root emission fter 90 to 120 d of the experiment (Ligrreto, 2011). FIGURE 4. Root presence in M. rupestris rnches t IBA tretments of 0 nd 1,000 mg L -1 (left to right). Photos: Veloz-Sun, C. nd S. Durán-Css. The results otined in the control tretment might e due to the influence of uds nd leves present in the lyered rnch, which stimulte rooting through the production of uxins in these orgns (Hrtmnn et l., 2010). In turn, rnch girdling cuses lockge in the cropetl nd sipetl flow of photossimiltes in the phloem, cusing n ccumultion of severl metolites (orgnic compounds, crohydrtes, uxins, etc.) ove this region (Alves-de Oliveir et l., 1999) tht fvors the formtion of cllus nd dventitious root primordi. Furthermore, the sence of light in the re of root formtion, use of sustrtes to provide continuous moisture, nd moderte tempertures re fctors tht fvor rooting in the girdling zone (Rmirez-Villloos nd Urdnet- Fernández, 2004; Hrtmnn et l., 2010). In this sense, the Durán-Css, Veloz-Sun, Mgnitskiy, nd Lncheros: Evlution of uv cmron (Mcleni rupestris Kunth A.C. Smith) propgtion... 21

5 results oserved in the control tretment might e due to the endogenous content of phytohormones in the rnches nd provision of eneficil conditions for rooting. However, the exogenous ppliction of uxin-type growth regultors cn speed up the rooting process (Hckett, 1988; Bcrín et l., 1994; Rhmn et l., 2000; Nz nd Aslm, 2003), which ws reflected in the emission of first roots t 60 d of the experiment for the tretments with IBA, with mximum length of 3.5 cm. Contrry to this, the control tretments presented only development of cllus for the sme period of oservtion. Numer nd length of dventitious roots The results show tht the tretments of 500 nd 1,000 mg L -1 IBA produced the highest numer of dventitious roots with lyering (Figs. 2 nd 3), with verge vlues of 27 nd 32 roots, respectively, which represent out two to three times higher root emission thn tht oserved in the control, which hd n verge of 12 roots. Similr results were found y Nz nd Aslm (2003) in mngo (Mngifer indic L.) ir lyers, where the tretments with 2,000 nd 3,000 mg L -1 IBA produced n verge of 39 nd 31 roots, respectively, while the control tretment hd only 25 roots. As shown in Fig. 2, there ws reduction of root emission in the 2,000 mg L -1 IBA tretment s compred with the other hormone tretments. However, sttisticlly significnt differences existed etween the IBA tretments nd the control. Likewise, ccording to the polynomil regression model, the highest numer of roots in the M. rupestris lyerings ws t pproximtely 1,200 mg L -1 IBA, so this IBA concentrtion could e the most suitle for this form of propgtion. In Fig. 4, one cn see tht n exogenous ppliction of uxin hd n effect on root formtion for the tretments of 0 mg L -1 (control) nd 1,000 mg L -1 IBA. This ehvior is consistent with the findings of Pul nd Aditi (2009), where the mximum numer of roots in ir lyers mde on Syzygium jvnic ws otined with the concentrtion of 1,000 mg L -1 IBA, while the concentrtions of 500 nd 2,500 mg L -1 showed no significnt differences compred to the control. In studies of Jtroph curcs L., the root numer incresed with the incresing concentrtion of npthtlenecetic cid (NAA) nd IBA to yield the mximum numer of roots in lyered rnches treted with 1,500 mg L -1 NAA followed y IBA t the sme concentrtion (Dhillon et l., 2011). Dhillon et l. (2011) showed tht the effect of growth regultors (indolecetic cid IAA, NAA, IBA) nd their concentrtions significntly influenced the rooting percentge nd numer of roots, ut did not significntly ffect the length of the roots of Jtroph curcs L. in ir lyers. This grees with the findings of the length of dventitious roots in lyers of M. rupestris, where no significnt differences etween the tretments were found; ll roots hd n verge length of 5.0 cm. According to Tchoundjeu et l. (2002), uxins promote meristemtic differentition, nd thus, contriute to the formtion of root primordi in the girdling zone of the rnch. For this reson, exogenous IBA might ffect the numer of dventitious roots more, nd to lesser extent the length of the roots. Weight of dventitious roots nd cllus Auxins promote the formtion of cllus nd dventitious root primordi, ecuse they stimulte cell division; for this reson the ccumultion of uxins ove nd elow the girdling zone fvors the formtion of roots nd cllus in this specific re (Tiz nd Zeiger, 2006). The cllus in the girdling zone of the rnches ws oserved s roundish mss of yellowish-white color, composed of presumly undifferentited cells (Fig. 3). According to Hrtmnn et l. (2010) nd Colodi et l. (2008), the cllus cells could further differentite nd form root primordi or serve s protective lyer for the formtion of dventitious roots from other tissues of the lyered rnch. In most plnts, cllus formtion is independent on dventitious root formtion, lthough the simultneous ppernce of these tissues my hppen ecuse oth re influenced y the sme environmentl fctors (Slisury nd Ross, 1994; Burch nd McGrw, 1996; Acost et l., 2000; Colodi et l., 2008). The plnts of the Ericles order differ in this feture nd cn form dventitious roots directly from cllus, such s te (Cmelli sinensis L.) plnts (Koyuncu nd Blt, 2004), or from cmium nd phloem, such s Vccinium sp. plnts (Mhlstede nd Wtson, 1952). However, in order to know whether the formtion of cllus is independent on dventitious root formtion in M. rupestris, histologicl study nd nlysis of the ntomicl chnges tht occur during the rooting of this species re needed. During the period of evlution, the dventitious roots were trnslucent in ppernce, ut in one specific cse, reddish roots were oserved growing out of the lck plstic; this might possily e due to the production of nthocynins in the dventitious roots exposed to light, which re the 22 Agron. Colom. 31(1) 2013

6 Dry weight (g) y=-8,78e-08x 2 +1,95E-04x+7,49E-02 R 2 =0,958 y=-1,33e-07x 2 +3,15E-04x+1,21E-01 R 2 =0, ,000 1,500 2,000 2,500 IBA concentrtion (mg L -1 ) Cllus Roots Fresh weight (g) y=-8e-07x 2 +0,002x+0,834 R 2 =0,865 y=-1e-06x 2 +0,002x+0,829 R 2 =0, ,000 1,500 2,000 2,500 IBA concentrtion (mg L -1 ) FIGURE 5. A. Dry weight of roots nd cllus t different IBA concentrtions. B. Fresh weight of roots nd cllus t different IBA concentrtions. Tretments with the sme letters re not significntly different ccording to Tukey test (P 0.05). The rs indicte stndrd devition. pigments tht ply n importnt role in photoprotection (Tiz nd Zeiger, 2006). Vriles of oth fresh nd dry weight of the roots nd cllus ehved similrly to the vrile of numer of roots, presenting the lowest vlues t 0 mg L -1, the highest vlues t 500 nd 1,000 mg L -1, nd decresed vlues t 2,000 mg L -1 IBA. For the tretments of 500 nd 1,000 mg L -1 IBA, dry weight of 0.14 nd 0.19 g for the roots nd 0.30 nd 0.28 g for the cllus, respectively, nd fresh weight of 1.48 nd 2.15 g for the roots nd 2.62 nd 2.70 g for the cllus, respectively, were recorded (Fig. 5). In generl, pplictions of 500 nd 1,000 mg L -1 IBA resulted in mssive cell growth (cllus + roots) in the cut re. According to Hrtmnn et l. (2010), cllus formtion is n event tht precedes rhizogenesis nd, therefore, it is ssumed tht n increse in cllus weight should e reflected in n increse of root weight (Fig. 5). However, the histologicl origin of dventitious root primordi in M. rupestris is unknown, nd there is lck of knowledge on whether cllus formtion is prerequisite for the formtion of the roots. Compring these vriles, the dry nd fresh weights of the cllus were high in ll IBA tretments, presenting sttisticlly significnt differences etween the IBA tretments nd control (Fig. 5). As for the root dry weight, no sttisticlly significnt differences etween the tretments were found (Fig. 5A), however, when compring the fresh weight of roots, the differences were sttisticlly significnt etween the tretments of 500 nd 1,000 mg L -1 IBA nd the control. According to Sánchez et l. (2009), in ir lyers performed on guv (Psidium gujv) plnts, with incresing concentrtions of IBA (2,000, 4,000, nd 6,000 mg L -1 ), cllus formtion ws reduced, nd its formtion ws completely inhiited t the concentrtion of 6,000 mg L -1 IBA. A similr trend ws otined in our study for M. rupestris, where concentrtions higher thn 1,500 mg L -1 IBA resulted in reduction of dry nd fresh weights of the roots nd cllus. This might e explined y the findings of Moreir et l. (2009), which show tht IBA in ir lyers during periods of ctive growth (high endogenous uxin levels) could rise hormone levels ove the optiml concentrtions, leding to decrese in rooting. Likewise, lthough IBA increses the elsticity of cell wlls, ccelerting division, excessive concentrtions of the hormone my inhiit this process (Rhmn et l., 2000). In the cse of propgtion y lyering in the vocdo (Perse sp.), the ppliction of IBA incresed the rooting rte, with concentrtion of 10,000 mg L -1 eing the most efficient, otining 74% rooting nd presenting the highest numer of roots y lyer compred to other tretments (Alves-de Oliveir et l., 1999). According to our findings, one should properly choose the dose of growth regultors ecuse the optiml concentrtions vry within the studied species (Kozlowski et l., 1991); for propgtion of M. rupestris with ir lyering, the concentrtion of 1,200 mg L -1 IBA produced the highest dry nd fresh root weights. Growth of lyered rnches The verge growth rte of the lyered rnches ws 1.49 cm per month, while the rnches without lyering presented growth of 2.78 cm per month (Fig. 6). Therefore, the Durán-Css, Veloz-Sun, Mgnitskiy, nd Lncheros: Evlution of uv cmron (Mcleni rupestris Kunth A.C. Smith) propgtion... 23

7 Longitudinl growth rte (cm/month) FIGURE 6. Monthly longitudinl verge growth rte of lyered nd nonlyered rnches. Tretments with the sme letters re not significntly different ccording to Tukey test (P 0.05). The rs indicte stndrd devition. lyering dversely ffected the longitudinl growth of the rnches due to phloem interruption cused y girdling, nd the ppliction of exogenous uxins did not completely counterct this effect. According to Ds et l. (1997), during the formtion of root primordi in ir lyers, grdul consumption of monoscchrides is generted. In turn, the stge of root development decreses the level of nitrogen in the rnches (Ds et l., 1997). The cictriztion of the wound cused y cutting, followed y consumption of photossimiltes nd nutrients for the formtion of cllus nd dventitious roots, might e the cuse of the decrese in growth of lyered rnches (Fig. 6), which could ffect future flowering nd fruiting in the sme. In compring the longitudinl growth of rnches treted with IBA, it my e oserved tht t concentrtions of 1,000 nd 2,000 mg L -1 IBA, the rnches hd the highest growth rtes, with vlues of 1.9 nd 2.1 cm per month, respectively. These two tretments hd sttisticlly significnt differences when compred with the tretments of 0 nd 500 mg L -1 IBA which ccounted for 1.1 nd 1.0 cm per month growth rtes, respectively (Fig. 6). The constnt contriution of endogenous uxins to supicl regions of the stem is required for the elongtion of their cells; dditionlly the exogenous uxins in cut sections of the stem stimulte their growth (Tiz nd Zeiger, 2006). This could help to explin why the tretments with higher IBA concentrtions hd the highest longitudinl growth rtes in lyered rnches of M. rupestris. Presumly, rnch with lrger numer of nodes nd leves in ctive growth would ring more endogenous uxins to the re ,000 2,000 Without lyering IBA concentrtion (mg L -1 ) of girdling; however, this vrile ws not evluted in our study ecuse ll rnches hd similr size t the time of lyering. For the dimeter growth rte (thickness) in the rnches, no significnt differences were found etween the lyered nd intct rnches; lso, no effect of the IBA ppliction on this vrile ws found. Both the thickness nd degree of lignifiction of the rnch s well s the rte of photosynthesis in the rnch could e indirect indictors of the mount of crohydrtes ville for the formtion of cllus nd dventitious roots, therefore it would e importnt to ssess the effect of these vriles on the lyering of M. rupestris in future studies. Conclusions nd recommendtions The ppliction of exogenous uxins, such s indoleutyric cid, to stimulte the formtion of dventitious roots on the lyered rnches of M. rupestris is fesile due to the fct tht the tretments with uxins promoted rooting s compred with the control. A study on the effect of the tretments on post-trnsplnt survivl of plnts, vegettive growth rte nd production is recommended. The results show tht concentrtion of pproximtely 1,200 mg L -1 IBA resulted in the lrgest numer of roots nd the highest dry weight of roots, nd could e considered s the optiml concentrtion for propgtion of M. rupestris with ir lyering. The propgtion of M. rupestris with ir lyers could e n effective method for consistently otining plnt mteril, either for the purposes of production or ecologicl restortion of pármo nd supármo res in Colomi. Acknowledgements The uthors re grteful to the stff of the Jose Celestino Mutis Botnicl Grden, Bogotá, for llowing this study on Mcleni rupestris plnts elonging to the pármo collection, prticulrly to Mr. Ricrdo Pcheco, Coordintor of the Sustinle Use Project, Scientific Office. Literture cited Acero, L. nd H. Bernl Guí pr el cultivo, provechmiento y conservción de l uv cmron: Mcleni rupestris (H.B.K) A.C. Convenio Andrés Bello y Áre de Cienci y Tecnologí No Bogot. Acost, E., M. Sánchez, nd M. Bñón Auxins. pp In: Azcón-Bieto, J. nd M. Tlón (eds.). Fundmentos de fisiologí vegetl. 2 nd ed. McGrw-Hill. Intermericn de Espñ, Brcelon, Spin. 24 Agron. Colom. 31(1) 2013

8 Adrince, G. nd F. Brison Propgtion of horticulturl plnts. 2 nd ed. McGrw-Hill, New York, NY. Aguilr, M. nd S. Torres Protocolo de uso y provechmiento de l uv de nís, Cvendishi rctet (Ruiz y Pvón ex Jume Sint. Hillire) Hoerold, en mtorrles ndinos del Altiplno Cundioycense. Instituto de Investigción de Recursos Biológicos Alexnder von Humoldt; MADR; Cámr de Comercio de Bogotá, Bogot. Alves-de Oliveir, A., O. Crlos-Koller, nd A. Villegs-Monter Propgción vegettiv de gucte selección 153 (Perse sp.) por codo en contenedor. Rev. Chpingo, Serie Hort. 5, Amy, N. nd H. Rincón Contriución l estudio de l uv cmron (Mcleni rupestris H.B.K.) y su posile utilizción en l industri limentici. Undergrdute thesis. Deprtment of Chemistry, Universidd Ncionl de Colomi, Bogot. Bcrín, M., M. Benincs, V. Andrde, nd F. Fereir Enrizmento de estcs éres de goieir (Psidium gujv L.): efeito do ácido indolutírico sore inicição rdiculr. Rev. Cient. 22, Burch, L. nd B. McGrw Citoquinins. pp In: Azcón-Bieto, J. nd M. 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9 Snow, D Tropicl frugivorous irds nd their food plnts: world survey. Biotropic 12(1), Sturm, H. nd J. Rngel Ecologí de los Pármos Andinos. Editor Gudlupe. Bogot. Tiz, L. nd E. Zeiger Fisiologí vegetl. Volumen II. 3 rd ed. Puliccions de l Universitt Jume I, Cstelló de l Pln, Spin. Tchoundjeu, Z., M. Avn, R. Lekey, A. Simons, E. Ash, B. Dugum, nd J. Bell Vegettive propgtion of Prunus fricn: effects of rooting medium, uxin concentrtions nd lef re. Agrofor. Systems. 54, Um, A., M. Prd, C. Ortiz, I. Alrcón, nd L. Rosero Dispersión de semills por ves frugívors en dos formciones vegetles del Prque Nturl Municipl Rncherí PNMR. Fculty of Sciences, Universidd Pedgógic y Tecnológic de Colomi (UPTC), Pip, Colomi. Vásquez, R. nd G. Coímr Fruts silvestres comestiles de Snt Cruz. 2 nd ed. Fundción Amigos de l Nturlez. Snt Cruz, Bolivi. 26 Agron. Colom. 31(1) 2013