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Agrociencia
versión On-line ISSN 2521-9766versión impresa ISSN 1405-3195
Agrociencia vol.50 no.7 Texcoco oct./nov. 2016
Natural renewable resources
Cedrela odorata L. mini-cutting rooting
1 Posgrado en Ciencias Forestales.
2 Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP). Campo Experimental El Palmar. 68410. Tezonapa, Veracruz, México.
3 Fitosanidad. Colegio de Postgraduados. Campus Montecillo. 56230. Carretera México-Texcoco, km. 36.5. Montecillo, Estado de México, México. (marcosjc@colpos.mx).
4 Centro Nacional de Recursos Genéticos (CNRG-INIFAP). 47600. Boulevard de la Biodiversidad #400. Tepatitlán, Jalisco, México.
A massive and low-cost vegetation propagation is required to establish clonal plantations of Cedrela odorata L. Developing appropriate protocols is necessary to fulfill all the requirements of the cutting rooting system. This study evaluated the effects of the substrate (perlite and a 2:1:1 peat, perlite, and vermiculite mix), the type of cutting (basal, intermediate, and apical) and the concentration of indole-butyric acid (IBA; 0, 1500, 5000, and 10 000 ppm) in the rooting of C. odorata cuttings. All cuttings tested were at least 25 cm in lenght, and 1 cm in diameter. The substrate type-IBA interaction will improve, in a different ratio, the production of adventitious roots according to the type of cutting. Experimental design was completely random, in a factorial arrangement, with ten replications. After 20 weeks, the cuttings showed an average rooting of 52.7 %. Perlite increased rooting over 50 %, with regard to the mix; meanwhile an IBA of 1500 ppm favored a 73.3 % rooting. The best response to rooting came from apical cuttings: the number and length of adventitious roots increased over 50 %, in comparison with the other two types of cuttings. The interactions between factors were not significant; however, apical cuttings with an IBA of 1500 ppm, rooted in perlite, was the better combination, with a 90 % rooting. Therefore, these are the best conditions for C. odorata rooting.
Key words: American cedar; Cedrela odorata; vegetative propagation; cutting; cloning
Para el establecimiento de plantaciones clonales de Cedrela odorata L. se requiere la propagación vegetativa masiva y económica. El sistema de enraizamiento de estacas cubre estos requerimientos, por lo que se necesita desarrollar protocolos apropiados. En este estudio se evaluaron los efectos del sustrato (agrolita y una mezcla de turba-agrolitavermiculita 2:1:1), el tipo de estaca (basales, intermedias y apicales) y la concentración de ácido indolbutírico (AIB; 0, 1500, 5000 y 10 000 ppm) en el enraizamiento de estacas de C. odorata. Todas las estacas probadas tuvieron longitud mayor a 25 cm y diámetro de 1 cm. La interacción tipo de sustrato y AIB favorecerá, en diferente proporción, la producción de raíces adventicias según el tipo de estaca. El diseño experimental fue completamente al azar, con arreglo factorial y diez repeticiones. Después de 20 semanas el enraizamiento promedio fue 52.7 % de estacas. La agrolita aumentó más de 50 % el enraizamiento, respecto a la mezcla, y 1500 ppm de AIB favorecieron el enraizamiento en 73.3 %. Las estacas apicales tuvieron respuesta mayor al enraizamiento, pues aumentaron más de 50 % el número y la longitud de raíces adventicias, respecto a los otros dos tipos de estacas. Las interacciones entre los factores no fueron significativas; pero la combinación mejor, con 90 % de enraizamiento, fue estacas apicales con 1500 ppm de AIB, enraizadas en agrolita. Así, estas condiciones son las mejores para el enraizamiento de C. odorata.
Palabras clave: Cedro rojo; Cedrela odorata; propagación vegetativa; esqueje; clonación
Introduction
Cutting rooting is the most important means of vegetative propagation and massive production. In trees, this allows the transference of all the genetic potential of the donor material and the creation of genetic uniformity in the resulting individuals. Subsequently, total genetic variance is captured and exploted and genetic profit is kept through genetic improvement (Zalesny et al., 2003). Additionally, this is a low-cost, relatively quick, and simple method (Capuana et al., 2000; Husen, 2004; Tarragó et al., 2005), and it contrasts with propagation in vitro, which requires more infrastructure and lab equipment investment, and special culture media and work protocols (particularly in the case of forest species). Cutting rooting consists of getting tree branches segments and keeping them in favorable environmental conditions, in order to induce rhizogenesis and organogenesis (Thanuja et al., 2002; Husen and Pal, 2003; Liu et al., 2012). In Chile, Spain, Portugal, China, Canada, Brazil, Australia, New Zealand, and the USA, this method is used to develop commercial clonal plantations of Pinus radiata D. Don, Pinus taeda L., Gmelina arborea Roxb., Populus, Platanus, Salix, and Eucalyptus (Bedon et al., 2011).
The rooting ability is directly related with the mother plant’s maturity stage, so the ability to induce secondary roots decreases with the age of the material to be propagated (Klein et al., 2000). In order to obtain juvenile buds, it is necessary to prune plants with greater rooting capacity. (Ruíz et al., 2005; Trobec et al., 2005; Castillo et al., 2013). Juvenile tissue has a better response to growth controllers than old and fully differentiated cells, probably because the genome zones that control organ formation are less repressed and can be freed with auxins treatments (Druege and Kadner, 2008; Klopotek et al., 2010; Denaxa et al., 2012).
The plants’ genetic factors and physiological state control root induction, and environmental factors establish their elongation (Thomas and Schiefelbein, 2002; Castrillón et al., 2008). The cutting rooting’s success depends on the species, the age of the donor plant, the branch section from which the cuttings are removed, the rooting medium, and the growth regulators used (Itoh et al., 2002; Klopotek et al., 2012).
Endogenous auxins -which are translocated from the apical meristem to the plant baseare one of the major physiological factors (Ruíz et al., 2005). In some species, this endogenous compounds are not enough to induce rooting, and exogenous auxins must be applied. Indole-butyric acid (IBA) is the most frequently used auxin. It is necessary to know the appropriate dose (Ruíz et al., 2005), which is related with the endogenous contents, and, at the same time, with the cutting’s original position in the plant.
Regarding environmental factors, the type of substrate that will provide support and adhesion to the roots must be taken into consideration. Root formation requires high amounts of oxygen; therefore, substrate must have the following characteristics: enough aeration, gas exchange, appropriate drainage, appropriate temperature and pH, and being pathogen free (Bonfil et al., 2007; Saranga and Cameron, 2007).
In Mexico, Cedrela odorata L. (American cedar) is one of the priority species, owing to its industrial importance (Sánchez et al., 2003; Ramírez et al., 2008; Quinto et al., 2009), and is one of the most commonly used species in commercial plantations (CONAFOR, 2015). However, Hypsipyla grandella (Zeller) can damage its growth and limit successful commercial plantations (Navarro et al., 2004). Trees capable of resisting this moth invasion (Sánchez et al., 2003) were detected and they are candidates for massive reproduction. Cutting rooting is a recommendable option for the multiplication of genotypes with superior features (Marrón et al., 2002; Kraiem et al., 2010); however, there are no protocols for the rooting of C. odorata cuttings.
Therefore, with the aim of developing a clonal propagation methodology through cutting rooting, an evaluation was performed about the effect of indole-butyric acid (IBA), the position of the cutting in the plant and the type of substrate in the rooting of juvenile cuttings of C. odorata, in humidity and temperature controlled conditions. Substrate typeIBA interaction will improve, in a different ratio, the production of adventitious roots according to the type of cutting.
Materials and methods
Localization and vegetal material
The seeds of the 99 family used for plant production were gathered on May 2012, at the seed orchard of Campo Experimental El Palmar, in Veracruz, which belongs to the Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias. The seeds were sowed on April 2013, in a greenhouse of Colegio de Postgraduados, in Texcoco, Estado de México. A 2:1:1 peat, perlite, and vermiculite mix was used as substrate; it was contained in 1 L. polyethylene forestry tray cells, with 10 g of controlled-release fertilizer per cell. Germination began on the 15th day, and was finished on the 35th day, with a 95 % growth. Subsequently, each month, frequent irrigation and fungicide (captan® 1 g L-1) were applied to the plant.
Mini-cutting production
Pruning at 6 and 12 months were carried out to stimulate the growth of juvenile material, 6 cm over plant collar. In order to evaluate the impact of the cutting’s position in the rooting of each bud, the apical, intermediate, and basal sections were separated. The selected buds were 25 cm in length, and 1 cm in diameter. The leaves of each twig were cut, and three 7 cm-long cuttings were obtained.
Substrate preparation
Substrate consisted of: perlite and 2:1:1 peat, perlite, and vermiculite mix. Both were sterilized for 40 minutes with water vapor; then, they were cooled and placed in a rooting chamber.
Establishing the experiment
Irrigation and temperature were controlled in a semiautomatic 2.7x1.2x0.7 meter rooting chamber. Irrigation was scheduled to distribute 2250 mL of distilled water in 10 daily irrigations. Relative humidity was kept over 80 % and temperature fluctuated between 18 and 25 °C during the day; when temperature reached 25 °C, an automatic fan cooled the chamber until temperature reached 18 °C.
As asepsis control, cuttings were placed in a 5 % Captán® solution during 15 min; then, to establish the best dose for rooting, cuttings were immersed in 1500, 5000 y 10 000 ppm of Radix® solutions, during 5 seconds; industrial talcum powder was used as control.
Substrates were humidified with distilled water. Cuttings were placed in 4 cm deep holes and with a 10x10 cm spacing, pressing the substratum around them, without damaging the tissues.
Experimental design and variables evaluation
This experiment had a completely random design, with a 2x3x4 factorial arrangement, as follows: two types of substrate, three cutting cut position, and four different IBA concentrations. Ten repetitions were established (10 cuttings per treatment, 240 in total). The experiment was carried out during the fall-winter season (August 26th 2014 January 13th 2015); the presence of pathogens was monitored by means of regular evaluation, and 20 weeks later the percentage of live cuttings, cuttings rooted by treatment, the number and length of primary and secondary roots per cutting were evaluated.
Prior to analyzing variance, the variable expressed as a percentage (Y) was transformed with the arccosine function of the square root of the original value expressed as a decimal fraction
where: Y ijkl is the response variable of i th type of cutting, with the j th IBA dose, in the k th type of substrate; μ is general average; E i is the fixed effect of i th type of cutting (apical, intermediate, and basal); D j is the fixed effect of the j th IBA dose (0, 1500, 5 000 y 10 000 ppm); S k is the fixed effect of the k th type of substrate (2:1:1 peat, perlite, and vermiculite mix vs. perlite); EDi j is the interaction between the i th type of cutting and the j th IBA dose; ES ik is the effect of the interaction between the i th type of cutting and the k th type of substrate; DS jk is the effect of the interaction between the j th IBA dose and the k th type of substratum; EDS ijk is the interaction between the i th type of cutting, the j th IBA dose, and the k th type of substrate; and, E ijkl is the random error related to Y ijkl observation.
Results and discussion
The effect of substrate, IBA dose, and type of cutting was significant (p≤0.05) in the cuttings’ survival, rooting and morphology. However, there was no effect in their interactions (Table 1).
Table 1 Probability values (p) in the variance analysis of rooting variables in Cedrela odorata L. cuttings.
Fuente de variación | Superv. (%) | Estacas enraizadas (%) | Número de raíces | Longitud de raíces | ||
Prim. | Secun. | Prim. | Secun. | |||
Sustrato | 0.0001† | 0.0001† | 0.4196 | 0.4462 | 0.0012† | 0.0749 |
Dosis de AIB | 0.0316† | 0.0454† | 0.0141† | 0.0079† | 0.0707 | 0.4370 |
Tipo de estaca | 0.0072† | 0.0453† | 0.0327† | 0.0073† | 0.0037† | 0.0894 |
Sustrato*Dosis | 0.9793 | 0.8502 | 0.8174 | 0.5978 | 0.0786 | 0.2898 |
Sustrato*Estaca | 0.6807 | 0.6850 | 0.1459 | 0.0841 | 0.1377 | 0.0043† |
Dosis*Estaca | 0.2164 | 0.0513 | 0.8958 | 0.9438 | 0.9267 | 0.6807 |
Sustrato*Dosis*Estaca | 0.7858 | 0.7998 | 0.3844 | 0.4060 | 0.1985 | 0.3051 |
†Significant differences (p≤0.05). Sur: survival, Prim: primary roots, Second: secondary roots.
Effect of the substrate
The highest survival and rooted cuttings percentage occurred with perlite (Table 2). Perlite increased rooting and survival, in comparison with the 2:1:1 peat, perlite, and vermiculite mixture, in a two to one ratio. Primary root length was greater with mixture than with perlite, in a two to one ratio.
Table 2 Average value and standard error for each factor and treatment in the Cedrela odorata L. cutting rooting (20 weeks).
Factor | Superv. (%) | Estacas enraizadas (%) | Número de raíces | Longitud de raíces (mm) | ||
Primarias | Secundarias | Primarias | Secundarias | |||
Sustratos | ||||||
Mezcla | 38.3±4.8 b | 32.5±4.7 b | 14.9±6.6 a | 11.8±1.7 | 684±66 a | 73±35 a |
Agrol. | 82.5±3.9 a | 79.1±4.1 a | 12.1±2.0 a | 10.2±1.5 | 347±68 b | 59±28 a |
CV (%) | 8.4 | 8.7 | 10.5 | 8.6 | 8.6 | 4.8 |
Dosis de AIB (ppm) | ||||||
0 | 66.6±3.8 b | 63.3±4.9 b | 4.8±3.46 b | 5.2±1.7 b | 356±237 | 62±23 a |
1500 | 75.0±3.6 a | 73.3±3.6 a | 14.2±2.7 ab | 11.5±3.8 ab | 640±205 | 71±39 a |
5000 | 48.3±9.3 c | 43.3±.4.7 c | 21.2±3.4 a | 16.4± 1.4 a | 538±178 | 67±32 a |
10 000 | 41.6±4.6 c | 31.6±4.8 c | 11.1±2.5 ab | 9.5±1. ab | 292±202 | 53±28 a |
CV(%) | 9.1 | 9.5 | 9.6 | 7.9 | 8.9 | 4.9 |
Tipo de estaca | ||||||
Apical | 68.7±4.5 a | 66.2±4.8 a | 21.3±5.5 a | 16.7±4.8 a | 705±149 a | 75±09 a |
Inter. | 55.0±5.3 b | 52.5±8.1 b | 8.9±4.5 b | 7.1±3.3 b | 390±165 b | 60±08 a |
Basal | 35.0±11.8 c | 32.5±4.8 c | 4.4±2.6 b | 5.3±2.8 b | 148±156 b | 50±16 a |
CV (%) | 9.1 | 9.5 | 9.1 | 7.2 | 8.0 | 4.7 |
Prom. | 56.7 | 52.7 | 12.5 | 10.4 | 4.55 | 0.63 |
Different letters in a column, for each factor, indicate statistically significant differences (p≤0.05). Superv: survival, mixture: 2:1:1 peat, perlite and vermiculite v/v, Agrol: perlite, ppm: parts per million, CV: coefficient of variation, Inter: intermediate, Prom: General average.
The best rooting with perlite is the result of its physical features, such as porosity, aeration, drainage, and pH (Saranga and Cameron, 2007). Bonfil et al. (2007), in order to keep moderate temperatures and to allow appropriate aeration, a substratum must provide constant humidity. Perlite has these features, which favour the rooting of: Pinus leiophylla Schiede ex Schltdl. et Cham. (Cuevas et al., 2015); Pinus pinaster Ait. (Majada et al., 2010); Pinus elliottii Engelm. var. elliottii Little & Dorman, Pinus caribea var. hondurensis Barr. & Golf. (Hunt et al., 2011), Juniperus scopulorum Sarg., and Thuja occidentalis L. (Bielenin, 2003).
Effect of the IBA dose
In comparison with control, a 1500 ppm IBA increased rooting by 10 %; higher doses decreased it, probably as a result of toxicity. According to Mesén et al. (1997) some forestry species -such as Eucalyptus, Cupressus lusitánica Mill., Alnus acuminata Kunth y Tectona grandis L. (Husen y Pal, 2003)do not tolerate concentrations higher than 2000 ppm. With a IBA of 1500 ppm, the number of primary and secondary roots was three times and twice than in the control group. The length of primary and secondary roots did not have a significant effect (p≤0.05) with IBA, in comparison with the control group. However, there was more than 30 % difference (p≤0.05) in cutting survival between the best and worst treatments.
With a 5000 ppm dose, the number of primary and secondary roots was five and three times higher (p≤0.05) than the control group. Once cuttings have established their roots, enhanced IBA doses increase the transportation of substances towards the cutting’s base, as a result of which more roots are developed (Mesén et al., 1997).
The lowest percentage of rooted cuttings was obtained with 10 000 ppm (Table 2), attributable to auxin intoxication. Cuttings without IBA showed two times more rooting than with 10 000 ppm; this indicates that cuttings have enough endogenous auxins to foster rooting.
Auxins foster adventitious root development in cutting rooting (Bielenin, 2003; Castrillón et al., 2008; Kumar et al., 2011). Castillo et al. (2013) suggest using IBA in species with difficult rooting. Rooted cuttings percentage showed significant differences (p≤0.05) between IBA doses. Auxins stimulated cambium activity and reserve mobilization towards the place where roots begin (Dhillon et al., 2011). IBA improves rooting in Pinus jaliscana Pérez de la Rosa (Aparicio et al., 2006); Viccinium meridionale Swartz (Castrillón et al., 2008); Taxus globosa Schltdl. (Muñoz et al., 2009); Olea europaea L. (Denaxa et al., 2012), and Gmelina arborea Roxb. (Ruíz et al., 2005).
Effect of the position of mini-cuttings
Survival and rooting percentages of apical cuttings were 20 and 50 % higher, respectively, than those of intermediate and basal cuttings (Table 2). Apical cuttings favored the amount of primary and secondary roots -twice and four times, respectively-, with regard to intermediate and basal cuttings. Additionally, the primary roots of apical cuttings were twice longer than intermediates (which was the second best). There were no differences with regard to the length of secondary roots.
Apical cutting showed the best rooting, perhaps because of some physiological or anatomical condition related to the presence of leaf buds. According to Hartmann et al. (2013), apical cuttings root better than lateral cuttings, because apical leaf buds produce plentiful auxins, and stimulate radical development (Moore, 1984). Ruíz et al. (2005) noted that the degree of juvenility, tissue lignification, reserves content, and, in particular, the level of endogenous phytohormones are the factors involved in enchancing the rooting capacity of Gmelina arborea apical cuttings.
In some cases, basal cuttings root better, like in Populus, where it doubles intermediate and apical cuttings (Zalesny et al., 2003). Cuevas et al. (2015) noted that the rooting of P. leiophylla basal cuttings is 3.5 more likely to succeed. The reason for this variation between species is unknown, and it is yet to be researched. In this study, although apical cuttings had the best rooting response, using basal cuttings to propagate C. odorata select clones has not been rejected. However, increasing the rooting of basal cuttings percentage requires constant experimentation; therefore, most of the material would be used to maximize plant production.
Effect of the interaction between factors
Only the substrate-type of cutting interaction showed a significant effect in the length of secondary roots (Table 1). The combination of apical cutting with substrate (2:1:1 peat, perlite, and vermiculite mix) favored longer secondary roots, 96±11 mm. Root length decreased gradually according to the type of cutting, from the longest (apical) to the shortest (basal); however, root length decrease was greater with mixture (intermediate position cuttings; Figure 1).
![](/img/revistas/agro/v50n7//1405-3195-agro-50-07-919-gf2.gif)
Figure 1 Interaction of cutting position and substrate in the length of secondary roots. Each point represents the average with the standard deviation.
The root length induced in cuttings can show the speed of rooting; the longest cuttings had less time to put down roots (Ruiz et al., 2005). In our study, the secondary roots of apical cuttings probably appeared first in the mixture’s substrate than in the perlite. This could prove advantageous, as it would decrease the time that the cuttings stay in the propagation unit, the contamination risks, and the maintenance costs. However, this substrate is still not recommended, because it is still not known if the length of cutting roots established in perlite can reach the size than those in the mixture, in one or two weeks.
These results show that there is a big potential for C. odorata vegetal propagation through cutting rooting. More research is necessary to determine if the greatest rooting percentage can be achieved in less time. Other IBA concentrations (around 1500 ppm), high quality clones, and apical sections could also be evaluated.
Conclusions
Perlite, like substrate, favors the rooting and survival of cuttings. A 1500 ppm IBA dose increases the rooting capacity of cuttings; however, a higher dose inhibit it. With regard to primary and secondary roots, the apical position of cuttings favors survival and rooting. Therefore, the best combination for rooting is the apical position with an 15 000 ppm IBA.
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Received: November 2015; Accepted: June 2016