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Revista Chapingo serie ciencias forestales y del ambiente

versión On-line ISSN 2007-4018versión impresa ISSN 2007-3828

Rev. Chapingo ser. cienc. for. ambient vol.21 no.1 Chapingo ene./abr. 2015 

Macropropagation of Erythrina americana in a greenhouse: a potential tool for seasonally dry tropical forest restoration


Macropropagación de Erythrina americana en invernadero: una herramienta potencial para la restauración de bosques tropicales estacionalmente secos


Tara C. Fehling-Fraser1; Eliane Ceccon2*


1 Facultad de Ciencias Biológicas, Universidad Autónoma del Estado de Morelos. Av. Universidad 1001, col. Chamilpa. C. P. 62209. Cuernavaca, Morelos. MÉXICO.

2 Centro Regional de Investigaciones Multidisciplinarias, Universidad Nacional Autónoma de México. Universidad s/n, Circuito 2, col. Chamilpa. C. P. 62210. Cuernavaca, Morelos. MÉXICO. Correo-e:, tel.: (777) 317 5299 ext. 302 (*Autora para correspondencia).


Received: April 25, 2014.
Accepted: March 18, 2015.



Deforestation in Mexico has made restoration an urgent requirement. Erythrina americana is a multipurpose tree, useful for seasonally dry tropical forest restoration; however, it is poorly studied. Macropropagation of this species is advantageous when compared with sexual reproduction, since it requires scarification to germinate. In this study, the effects of the exogenous application of the phytohormone (indole-3-butyric acid) in the cuttings and/or a slow-release phosphate fertilizer (SRPF) in the substrate were evaluated on the growth and survival of E. americana cuttings in a greenhouse. We used a randomized experimental design of four blocks with 10 cuttings per treatment. We harvested 10 cuttings per treatment after 36, 66, 96 and 126 days of growth, and evaluated survival, total dry biomass production (TDB), relative growth rate (RGR), and resource allocation (RA). Cutting survival was high, averaging 95 %. Treatments did not affect total TDB, RGR or RA. However, the application of SRPF in the substrate showed significantly higher (P < 0.05) dry root biomass values. Growth periods also affect the RGR and RA. Macropropagation of E. americana was successful and non-labor-intensive, making this technique a feasible alternative for restoration projects mainly in low-resource rural communities.

Keywords: Cuttings, indole-3-butyric acid, slow-release phosphate fertilizer, relative growth rate, resource allocation.



La deforestación en México ha hecho que la restauración sea una necesidad urgente. Erythrina americana es un árbol multipropósito, útil para la restauración de bosques tropicales estacionalmente secos (BTES), pero es poco estudiado. La macropropagación de esta especie es ventajosa comparada con la reproducción sexual que requiere de semillas escarificadas para germinar. En este estudio se evaluaron los efectos de la aplicación exógena de la fitohormona ácido indol-3-butírico en esquejes y del fertilizante fosfatado de liberación lenta (FLL) en el sustrato, sobre el crecimiento de E. americana en invernadero. El experimento tuvo un diseño de cuatro bloques al azar. Diez esquejes se cosecharon por tratamiento después de 36, 66, 96 y 126 días de crecimiento. La sobrevivencia, biomasa seca total (BST), tasa de crecimiento relativo (TCR) y la asignación de recursos (AR) se evaluaron. La sobrevivencia promedio de los esquejes fue alta (95 %). Los tratamientos no afectaron la BST, TCR y AR; sin embargo, la aplicación de FLL en el sustrato mostró valores significativamente (P < 0.05) mayores de biomasa seca radical. Los periodos de crecimiento afectaron la TCR y AR. La macropropagación de E. americana fue exitosa y no requirió cuidados intensivos, por lo que esta técnica puede ser una alternativa viable para proyectos de restauración, principalmente en comunidades con bajos recursos.

Palabras clave: Esquejes, ácido indol-3-butírico, fertilizante fosfatado de liberación lenta, tasa de crecimiento relativo, asignación de recursos.



One of the most challenging problems facing the Mexican environment is deforestation. In 2010, the primary forest cover percentage in Mexico was only 18 % (Food and Agriculture Organization of the United Nations [FAO], 2010). Unfortunately, loss of forest cover is often neglected as it is commonly regarded as a sign of progress, which has led to the emergence of large portions of extremely degraded lands and unproductive farms (Ceccon, 2008).

Rapidly restoring these degraded areas or landscapes is not an easy task for researchers and lawmakers due to current ecological and socio-economic limitations. It is therefore urgent to develop strategies for the restoration of ecosystem services mainly based on the needs of local peasants (Aronson, Milton, & Blignaut, 2007; Ceccon, 2013). The seasonally dry tropical forest (SDTF) is one of the most prevalent biomes in the tropics where degradation is most problematic due to the SDTF's easy conversion to exploitation (Ceccon, Huante, & Rincon, 2006), with extensive areas of these forests having been severely disturbed by frequent forest fires, cattle raising and wood extraction (Sánchez-Azofeifa & Portillo-Quintero, 2011). In Mexico, Trejo and Dirzo (2000) estimated that only 27 % of the original SDTF cover remained as intact forest, representing about 3.7 % of the total area of the country. However, most restoration studies have focused on the propagation of temperate or moist tropical species, while little is known about Mexican SDTF restoration (Bonfil & Trejo, 2010; Castellanos-Castro & Bonfil, 2013; Cervantes, Arriaga, Meave, & Carabias, 1998; Cervantes, López, Salas, & Hernández, 2001).

Selection of species employed in restoration or reforestation projects depends mainly on the available scientific information that is useful in facilitating their propagation and management (Cervantes et al., 2001). Furthermore, some useful species present difficulties to germinate because they exhibit dormancy and so they require seed scarification; which may be a problem for their reproduction in small places that lack appropriate technology (Bewley & Black, 1994; Bonfil-Sanders, Cajero-Lázaro, & Evans, 2008). Although macropropagation is one of the main methods currently used in the multiplication of species in modern fruit orchards, this technique has been implemented with only a few native species for restoration purposes (Souza & Araújo, 1999; Castellanos-Castro & Bonfil, 2013). Given the commercial success of vegetative propagation, it is important to evaluate this technique with native species, focusing on restoration goals. This in turn demands a deep understanding of the growth eco-physiology of the species centered, in particular, on the feasibility of its cultivation under nursery conditions (Bonfil & Trejo, 2010; Cervantes et al., 1998).

Erythrina americana Miller, belonging to the Leguminosae family, is a native species from the Mexican seasonally dry tropical forest (Comisión Nacional para el Conocimiento y Uso de la Biodiversidad [CONABIO], 2009). This species is well known and it has a cultural importance for local peasants because of its multipurpose benefits: it is used in living fences, it provides shade and organic matter to crops and it is widely used as an ornamental tree (Flores, 2002). Its leaves have a high nutritional value and the tree is resistant to repeated pruning (Flores, 2002). Erythrina americana flowers are highly valued as an ingredient for various traditional dishes (García-Mateos, Soto-Hernández, & Vibrans, 2001) and the seeds and wood are used in indigenous handcrafts (Niembro, 1992). In traditional Mexican medicine, different parts of the plant are used for their attributable antidotal, narcotic, laxative, diuretic, expectorant, anti-inflammatory, sedative, anti-asthmatic, anti-malarial and anti-dermatitis properties (Argueta, Cano, & Rodarte 1994; García-Mateos et al., 2001). Another advantage of E. americana is that it is a nitrogen-fixing species establishing symbiosis with rhizobiums of the "caupi" group, which gives it a high potential for improving soil fertility and accelerates the regeneration of other species (García-Mateos et al., 2001). In fact, Suárez and Equihua (2009) mentioned the great potential of E. americana for the restoration of degraded soils. This species is also one of the species recommended for restoration by the governmental Program for Sustainable Use of Natural Resources of Morelos State in Mexico (Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias-Secretaría de Agricultura, Ganadería, Desarrollo Rural, Pesca y Alimentación [INIFAP-SAGARPA], 2014).

Macropropagation of E. americana is especially advantageous over sexual reproduction because its seeds exhibit dormancy due to the presence of a hard and impermeable coat, which requires physical or chemical scarification (Argueta et al., 1994), hindering its management in nurseries. In macropropagation, auxins are often considered a main agent responsible for inducing adventitial rooting, which is mostly useful in species that have a difficult rooting. Among auxins, the most widely used substance in rooting cuttings that has shown the best results for most forest species is indole-3-butyric acid (IBA) (Cunha, Wendling, & Júnior, 2008; Valmorbida, Boaro, Lessa, & Salerno, 2008). On the other hand, slow-releasing phosphate fertilizers have been largely used in agriculture but little is known about their effects on cuttings of forest species in dibble-tube nurseries (Rose, Haase, & Arellano, 2004). In this context, this study evaluates the effect of applying indole-3-butyric acid (IBA) in E. americana cuttings and/ or slow-release of phosphate fertilizer to the substrate, on the survival, growth and resource allocation of this species in greenhouse conditions using destructive samples, in a way that would be feasible for restoration projects in seasonally dry tropical forests in Mexico.



Study area

We collected the cuttings from Buenavista del Monte (18° 56' 41'' N and 99° 18' 33'' W), Morelos, a community with high levels of environmental degradation. The climate is semi-warm with an average monthly temperature of 21 °C to 24 °C, and annual seasonal precipitation of 1,000 to 1,200 mm, most of which falls between June and October (Comisión Nacional del Agua [CONAGUA], 2007; Instituto Nacional de Estadística y Geografía [INEGI], 2000). The vegetation is represented by deciduous and perennial species of tropical deciduous forests (Gómez-Garzón, 2002).

Greenhouse environment: The cuttings were cared for under similar environmental conditions to the study site and monitored by a digital device (HOBO Data Logger, model H21-002, Onset, USA); the average light intensity was 258.6 uE, temperature 21.7 °C and soil humidity 0.049 θ.

Study design

A total of 160 cuttings averaging 15 cm in length and 10 g in weight, with at least three buds, were collected from the young lower branches of different trees of E. americana, and were subjected to the following treatments: 1) Exogenous phytohormone application of 0.3 % indolbutyric acid (IBA; Radix T 3000) on the cuttings, 2) Slow-release phosphate fertilizer (SRPF; triple superphosphate 0-45-0) in the substrate, 3) Exogenous phytohormone + SRPF, 4) Control. Four blocks with 10 cuttings per treatment were randomized in the greenhouse (in total 160 cuttings). The cuttings were placed in the polypropylene tubes of the dibble-tube system with peat-moss, vermiculite and perlite in a 7:1.5:1.5 proportion, under greenhouse conditions for four months. All the cuttings were uniformly fertirrigated with a N-P-K (9:35:7) commercial fertilizer (Western Fol 66 Plus) in a solution of 1 g·liter-1 (at 30, 40 and 50 days of growth) which subsequently increased to 5 g·liter-1 (from day 60, 70, 80, 90, 100, 110 to 120) due to root presence. The cuttings were manually watered daily.

To evaluate the effect of the treatments, 10 randomly selected individuals per treatment were destructively harvested at 36, 66, 96 and 126 days of growth. Each plant was separated into roots, shoots and leaves; and was then oven-dried at 80 °C for 6 to 8 h. We measured the following parameters: total dry mass, root, shoot and leaf dry mass and root length. These data were used to calculate the average survival percentage, total dry biomass production for each structure (root + shoot + leaves), and daily relative growth rate (RGR; calculated by fitting total dry-mass measurements to a Hunt and Parsons model [Hunt & Parsons, 1974]). Resource allocation over time was also calculated using the relationship between the root/shoot weight ratio (R/S), and the root length/dry weight ratio (RL/RDW). The analysis of these last variables started after 66 days because until this time cuttings did not exhibit roots yet, and the RGR was relative to day 36. To evaluate the average survival percentage in each treatment and in each growth period, cutting death was considered to have occurred when the cutting lacked roots and leaves, since some species tend to sprout before developing roots (Zahawi, 2005).

Statistical analysis

A one-way ANOVA was used for each harvest period to compare total dry biomass and dry biomass of shoots and roots among treatments. To evaluate the effect of treatments in the R/S and RL/RDW ratios and in the RGR, a factorial ANOVA analysis (four treatments x three growth periods) was used. Tukey tests (P ≤ 0.05) were used for the post hoc comparison of means.



The exogenous phytohormones and the slow-release phosphate applications were found not to affect either E. americana survival, which was high (around 95 %) (Figure 1), or total dry biomass production in the greenhouse in all growth periods (36 days, F = 1.980; 66 days, F = 1.665; 96 days, F = 0.516; 126 days, F = 0.794; P ≥ 0.05) (Figure 2a). Despite the important role of exogenous phytohormones in the growth of many tree species (Valmorbida et al., 2008), some recent studies have also found that exogenous phytohormones did not influence the development of tropical tree species (Baul, Mezbahuddin, Hossain, & Mohiuddin, 2010; Cézar et al. 2009; Leandro & Yuyama, 2008). On the other hand, in the last growing period (126 days) of this study, the cuttings fertilized with SRPF showed significantly higher dry root biomass (66 days, F = 1.254; 96 days F = 1.573; P ≥ 0.05; 126 days, F = 3.002; P = 0.047) than those that received exogenous phytohormones alone, as well as the control group, which showed similar results (Figure 2b). Since fertilization is a routine and necessary step in the dibble-tube cultivation system, the application of SRPF does not represent an extra investment of time or effort, making it an economical and effective technique to increase root biomass. A large root biomass before transplantation may improve the long-term survival of cuttings in degraded areas, especially in seasonal climate regions where this characteristic may allow plants to access the deeper, moister layers of the soil during the dry season (Ceccon et al., 2006). On the other hand, in the second harvest period (at 66 days of growth), the biomass production of the shoot structures resulted in a significant increase (36 days, F = 0.130; 96 days F = 0.464, 126 days, F = 1.516; P ≥ 0.05; 66 days, F = 6.777; P = 0.001) in the absence of SRPF (Figure 2c). It is possible that this is related to the low root biomass recorded during this period; thus, the effect of SRPF was indiscernible.

The exogenous phytohormones and the slow-release phosphate applications did not affect significantly the relative growth rate, root/shoot weight ratio or root length/root dry weight ratio in the different growth periods nor the interaction between these treatments and growth periods. However, these variables presented significant changes over the growth periods (Table 1).

In the last growth period (96-126 days), the cuttings presented a significantly higher RGR than in previous periods (Figure 3a), probably due to the large root proportion (4.3 % of total biomass), which increased the plant's ability to take advantage of the applied nutrients and water (Cairns, Brown, Helmer, & Baumgardner, 1997). This also indicates the appropriateness of maintaining the E. americana cuttings in the greenhouse for at least four months before transplanting them to the field in order to increase their survival probability in the dry season. In the present study, the RGR average of E. americana cuttings at 126 days was 1.92 g·g-1·day1 (Figure 3a), which was much higher than that of other fast-growing species of STDF reproduced by seeds: 0.097 g·g-1·day1 and 0.126 g·g-1·day1 as found by Huante et al. (1995) and Huante, Ceccon, Orozco, Sánchez, and Acosta (2012), respectively. According to Holl, Loik, Lin, and Samuels (2000), pasture is one of the most limiting barriers for restoration due to competition with seedlings, and one successful strategy is to establish forest recovery with high RGR species. Therefore, the high RGR of E. americana cuttings may be advantageous when compared to sexual reproduction for restoration of degraded pasture areas.

The R/S increase of E. americana cuttings over time (Table 1, Figure 3b) agrees with results obtained from other STDF species reproduced from seeds in greenhouse conditions (Ceccon, Almazo-Rogel, Martínez-Romero, & Toledo, 2012; Cervantes et al., 1998). The higher root/shoot weight ratio found in STDF species is an adaptive response to the severe seasonal conditions of low-moisture soils (Bullock, 1990; Martínez-Yrizar et al., 1992).

The reduction of the RL/RDW ratio over the time period (Table 1, Figure 3c) was probably due to the fact that as time goes by, the roots become thicker (due to increasing resource allocation) and longer. This root morphology not only gives this species a better ability to adapt in areas with low water availability (typical of SDTF during four to six months per year), but also provides it with certain benefits to successfully tolerate areas with low fertility (Ceccon et al., 2006). Larger roots are more capable of absorbing water and soil nutrients (Ceccon et al., 2006).

The foregoing results, taken together, suggest that macropropagation of E. americana is an important tool for SDTF restoration, particularly in low-resource rural communities, since it needs few external inputs in the greenhouse which reduces cost and labor.



The exogenous phytohormones and the slow-release phosphate applications did not affect the total dry biomass production or high survival of E. americana cuttings in the greenhouse in all growth periods. The application of slow-release phosphate in the substrate increased the root biomass of E. americana cuttings and could provide more plasticity and ability to adapt to the changing moisture conditions and to survive the long SDTF drought season, which is decisive for the planning and effectiveness of restoration projects. The relative growth rate and root/shoot dry weight ratio increased significantly while the root length/root dry weight ratio was reduced over the growth periods. Thus, the time lapse in the greenhouse proves to be a determinant variable in the macropropagation of E. americana. Cuttings should be maintained in the greenhouse for at least four months before transplanting to the field at the beginning of the rainy season. In this way, cuttings will be larger and the roots will present a higher biomass. The vegetative macropropagation of E. americana could be an important tool for SDTF restoration, mainly in low-resource rural communities since it requires few external inputs in the greenhouse, thereby reducing costs and labor.



We very much appreciate PAPIIT/UNAM grants IN300112 and IN300615, and Lynna Kiere for her useful comments.



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