Morphoanatomical aspects of stem cuttings of Selenicereus setaceus (Salm-Dyck) A. Berger ex Werderm. (Cactaceae)

Aspectos morfoanatômicos da estaquia de Selenicereus setaceus (Salm-Dyck) A. Berger ex Werderm. (Cactaceae)

Abstract

The cerrado pitaya (Selenicereus setaceus) is a Cactaceae with strong potential in the fruit market. For that reason, propagation cutting is an interesting alternative for fixing desirable characteristics for the species. Therefore, this work aimed to investigate the origin and morphoanatomy of adventitious roots from this species’ cuttings, an essential step to develop alternatives to its management. The first experiment was carried out in spring and tested peat-carbonized pine bark and peat-expanded perlite as substrates. The second was carried out in autumn, and tested three substrates: vermiculite-carbonized rice husk, sand and bio-stabilized pine bark, vermiculite, charcoal mill and additives. The following variables were evaluated: rooting (%), number of roots per cutting, length of the three largest roots (cm) per cutting, survival (%) and sprouting (%). Fresh samples were cross-sectioned and stained with toluidine blue for the anatomical analyses. In spring, the sprouting and the number of roots were higher when the peat-expanded perlite was used. In autumn, the substrate used had no significant influence on the number of roots, only on the rooting percentage and the mean root length. In addition, in spring, the number and mean length of the roots were also higher when the peat-expanded perlite substrate was used compared to autumn. Thus, it is not recommended to prepare cuttings in autumn, as there is no sprouting. In the pith and the cortex, mucilaginous cavities and starch are abundant, ensuring water and energy supply, which probably influence the species’ excellent rooting capacity. The formation of adventitious roots occurs from the interfascicular cambium, and these cross the perivascular fibres, cortical bundles, and the collenchyma, which, therefore, do not act as anatomical barriers. In regions where the periderm has already replaced the epidermis, there is no adventitious root formation, indicating that this tissue is a barrier to rooting. Although this species is recognized for its easy rooting, the type of substrate and the season influenced the variables assessed here.

Keywords:
plant anatomy; cactus; stem; propagation

Resumo

A pitaya do cerrado (Selenicereus setaceus) é uma Cactaceae que tem forte potencial no mercado de frutas. Portanto, a estaquia é uma alternativa interessante na fixação de caracteres desejáveis para a espécie. Diante disso, o objetivo do trabalho foi entender o processo de origem e formação das raízes adventícias a partir da estaquia da espécie, sendo um passo importante para a compreensão e desenvolvimento de alternativas ao seu manejo. Foram realizados dois experimentos: o primeiro na primavera, testando-se dois substratos: (turfa + casca de pinus carbonizada 1:1) e (turfa + perlita expandida 1:1), e o segundo no outono, testando-se três substratos: (vermiculita + casca de arroz carbonizada), areia e (casca de pinus bio-estabilizada + vermiculita + moinha de carvão vegetal + aditivos). Em ambos, avaliou-se as variáveis enraizamento (%), número de raízes por estaca, comprimento das 3 maiores raízes (cm)/estaca, sobrevivência (%) e brotação (%). Para as análises anatômicas, amostras frescas foram seccionadas transversalmente e coradas com azul de toluidina. Na primavera, tanto a brotação quanto o número de raízes foram maiores quando se utilizou o substrato turfa + perlita expandida (1:1). No outono, não houve influência significativa do substrato utilizado no número de raízes, mas sim na porcentagem de enraizamento e no comprimento médio das raízes. Em relação à época do ano, não se recomenda a confecção de estacas no outono, pois não há brotação. Além disso, na primavera, as variáveis número e comprimento médio das raízes quando se utilizou o substrato turfa + perlita expandida (1:1), também foram maiores quando comparadas ao outono. Tanto na medula, quanto no córtex, há uma grande quantidade de cavidades mucilaginosas e amido, garantindo um suprimento de água e energia, condições que provavelmente influenciam a grande capacidade de enraizamento da espécie. A formação de raízes adventícias ocorre a partir do câmbio interfascicular, e essas atravessam as fibras perivasculares, feixes corticais e o colênquima que, portanto, não atuam como barreiras anatômicas. Nas regiões onde a epiderme já foi substituída pela periderme, não há formação de raízes adventícias, o que indica esse tecido como uma barreira ao enraizamento. Mesmo sendo reconhecida pelo fácil enraizamento, o tipo de substrato e a época do ano influenciaram nas variáveis aqui testadas.

Palavras-chave:
anatomia vegetal; cacto; caule; propagação

1. Introduction

Pitaya is the generic name assigned to all fruits of hemiepiphytic columnar cacti belonging to the tribe Hylocereeae, subfamily Cactoideae, Cactaceae family, grouped into eight genera, among them: Selenicereus (A. Berger) Britton and Rose (Zappi and Taylor, 2020). In their natural habitat, pitayas develop on rocky massifs, tree trunks and sandy soils of high-altitude cerrado (Junqueira et al., 2002). Naturally, as they are succulent and adapted to arid conditions, pitayas are not plants that need fertile substrates or large amounts of water to complete their life cycle, which makes these species promising for cultivation anywhere in the national territory (Santos et al., 2022).

Currently, pitayas have been standing out in the fruit market for the diversity of colours of the epicarp and endocarp (Jalgaonkar et al., 2020). In addition, as a plant with potential for low-cost production and fruits marketed at a high value, pitayas are also good alternatives for small producers (Pires and Krauze, 2020). Among the species currently cultivated, Selenicereus setaceus (Salm-Dyck) A. Berger ex Werderm, known as pitaya-do-cerrado or “Saborosa”, stands out for being one of the four most commercialized pitaya species in Brazil (Santos et al., 2022).

In Brazil, S. setaceus occurs naturally in the states of Pará, Alagoas, Bahia, Ceará, Maranhão, Paraíba, Pernambuco, Piauí, Sergipe, Distrito Federal, Goiás, Mato Grosso do Sul, Mato Grosso, Espírito Santo, Minas Gerais, Rio de Janeiro, São Paulo, and Paraná (Zappi and Taylor, 2024). Therefore, it is possible to obtain matrices for seedling production locally anywhere in the national territory, which is also a factor that can reduce production costs and indicates adaptability to different edaphoclimatic conditions (Faleiro and Junqueira, 2021).

Pitaya production, recognized for its rusticity, was predominantly extractivist until the 1990s (Bastos et al., 2006). However, over the past two decades, growing consumer demand for exotic-looking fruits has driven the expansion of pitaya cultivation in Brazil. Today, Brazil ranks among the ten largest producers of this fruit worldwide (Mercado-Silva, 2018; Santos et al., 2022). Environmentally, pitaya cultivation offers a sustainable way to repurpose degraded areas without deforestation. Farmers can also extract honey from the bees that pollinate the plant’s flowers and repurpose the pruned stems as animal feed (Hartmann et al., 2002; IDAM, 2023). Pitaya has significant potential for use in the production of dyes, jellies, ice cream, sweets, yoghurts, and cookies. Additionally, it can be utilized in the cosmetics industry for products such as moisturizing creams, ointments, and lipsticks (Lima et al., 2021).

Pitaya cultivation is notably common in family backyards, as the plant adapts well to small spaces and is undemanding concerning nutrients and irrigation, presenting rapid growth (Santos et al., 2022). In just 100 square meters, 11 plants can be cultivated, yielding up to 275 fruits per year (Coelho and Oliveira, 2023). On a commercial scale, one hectare can accommodate 1,100 plants, with an annual production potential of approximately 10,400 kg (Santos et al., 2022). In Brazil, there are initiatives to select cultivars with thinner fruit spines, which would reduce the post-harvest cost (Junqueira et al., 2002). In addition, pitaya cultivars with a sweeter endocarp have also been the subject of research to diversify the use of the pulp recognized for its mild flavour (EMBRAPA, 2023).

Although pitaya can be propagated via seeds (Zee et al., 2004; Le Bellec et al., 2006), seminiferous propagation is not ideal for achieving uniformity in traits such as precocity and the fixation of selected genotypes. These traits are crucial for meeting the demands of producers and the consumer market, including fruit appearance, taste, and reduced production time (Hartmann et al., 2002; Gunasena et al., 2007). In addition, as a climbing plant in nature, pitaya naturally emits adventitious roots along the stem to attach the plant to the support where it climbs (Soffiatti and Rowe, 2020; Faleiro and Junqueira, 2021). Thus, it has a very promising characteristic for a species intended to propagate through cutting.

While studies have demonstrated the strong rooting potential of various pitaya species (Santos et al., 2010; Nascimento et al., 2018; Lone and Takahashi, 2019), information on the cutting propagation of Selenicereus setaceus remains limited. Furthermore, no anatomical studies in the literature detail the origin of adventitious roots in this genus or identify potential anatomical barriers that could reduce rooting capacity. As the species is easily rooted, we tested the hypothesis that the type of substrate used and the season will not harm the rooting of the species. Therefore, the objective of the present study was to evaluate the propagation potential of S. setaceus as well as the origin of its roots, using different substrates in propagules collected in two seasons of the year.

2. Material and Methods

The stems of S. setaceus were obtained from a matrix plant collected in the municipality of Armação dos Búzios, RJ, Brazil. The matrix is under cultivation in a pot at the Laboratory of Plant Anatomy and Biomechanics (LABV) of the Federal University of Paraná in the municipality of Curitiba, Paraná, Brazil. The species’ identified was confirmed by a specialist in the Cactaceae family. The cuttings were 10 cm long and packed in trays containing compartments with 65 cm³ of volume each. They remained on the LABV’s premises.

Two experiments were carried out (E1 and E2) in which different substrates were tested to evaluate the following variables: percentage of rooted cuttings (PR), number of roots per cutting (NRC), length of the three largest roots (MRL), percentage of sprouting and percentage of survival (Pereira et al., 2015).

Each experiment was completely randomized, with three replicates of 10 experimental units for each substrate used. First, the results were submitted for analysis of variance. Initially, the variances of the treatments were evaluated for their homogeneity by Bartlett’s test. The variables whose variances were homogeneous had the effects of the treatments evaluated by Tukey’s test at the 5% probability level. The software Assistat (Silva and Azevedo, 2002) was used to perform all statistical analyses.

The density values ​​of each substrate used in experiments E1 and E2 are found in Table 1.

Table 1
Dry mass density values ​​for each substrate used in the study.

Substrate	Density (kg/m3)
Peat + carbonized pine bark	65
Peat + expanded perlite	110
Carbonized rice husk + vermiculite	77
Sand	1700
Vivatto Plus®	240

• Source: Terral^®, Favorita Substratos^®, Global Minerio^®, Areal Costa^®, Vivatto Plus^®.

2.1. Experiment 1 (E1)

In the first experiment, conducted on 15th December 2023 (spring), commercial substrates Terral (a 1:1 mixture of carbonized peat and pine bark) and Sphagnotec (a 1:1 mixture of peat and expanded perlite) were tested. Sixty cuttings were made, 30 for each substrate. Each cutting received 40 mL of water twice a week. After 30 days of installation, the experiment was evaluated.

2.2. Experiment 2 (E2)

On 26th April 2024, E2 was installed (autumn) and was conducted in a completely randomized design with three substrates: vermiculite and carbonized rice husk (1:1), sand and Vivatto Plus substrate (bio-stabilized pine bark, vermiculite, charcoal mill, and additives). Each cutting received 40 mL of water every four days. The evaluation of the experiment occurred 66 days after its installation.

2.3. Anatomical analysis

The rooted cuttings from the treatment with vermiculite + carbonized rice husk were submitted to anatomical analysis at LABV. The fresh samples were cross-sectioned with 120 µm thickness using a vibrating blade microtome (5100mz Vibratome, Campden Instruments). The obtained serial sections were stained with 0.05% toluidine blue pH 6.8 (Feder and O’Brien, 1968) and mounted on slides with distilled water. Part of the samples were fixed in FAA50%, embedded in paraffin, sectioned in a rotating microtome and subjected to double staining. Lugol's dye was used to detect starch (Johansen, 1940) and Sudan III was used to detect cuticle (Sass, 1951). For the detection of collenchymal hypodermis, ruthenium red was used (Johansen, 1940). The images of the slides were obtained at the Center for Advanced Fluorescence Technologies (CTAF), at the Biological Sciences Sector of the Federal University of Paraná (UFPR) in Curitiba, Brazil, using a light microscope (Olympus BX51) with a digital camera attached.

3. Results

3.1. Experiment 1 (E1)

The highest percentage of rooting was verified in cuttings grown on peat and carbonized pine bark, reaching 83% of rooted cuttings (Table 2) and 17% of cuttings were alive but not rooted (Figure 1A). The treatments significantly differed in the mean root length and number of roots per cutting. Peat and expanded perlite provided a higher number of roots and longer ones than peat and carbonized pine bark (Figure 1B-C).

Table 2
Results of the percentage of rooted cuttings (PC), number of roots per cutting (NRC), mean root length (MRL) and sprouting (SP) of Selenicereus setaceus in spring and autumn.

Substrate	PR (%)	NRC	MRL (cm)	SP (%)
Spring
Peat + carbonized pine bark	83.00 a	2.70 b	0.64 b	0 b
Peat + expanded perlite	77.00 a	5.20 a	5.83 a	33 a
CV (%)	84.00	91.00	96.00	1.41
Autumn
Carbonized rice husk + vermiculite	66.66 b	2.23 a	3.32 a	0 b
Sand	73.33 b	2.97 a	1.53 b	0 b
Vivatto Plus®	90.00 a	2.90 a	1.54 b	0 b
CV (%)	27.42	42.70	30.39	-

• Means followed by different letters differ from each other at 5% significance by Tukey’s test. CV: Coefficient of variation.

Figure 1
Morphological aspectos of pitaya-do-cerrado (Selenicereus setaceus) rooting: (A-C) Cuttings 30 days after installation of Experiment 1. (D-F) Cuttings 66 days after installation of Experiment 2. (A) Unrooted cutting submitted to peat and carbonized pine bark; (B) Cuttings submitted to peat and carbonized pine bark; (C) Peat and expanded perlite; (D) Carbonized rice husk and vermiculite; (E) Sand; (F) Vivatto Plus.

The mean number of roots per cutting for the treatment with peat and carbonized pine bark was 2.70, while for peat and expanded perlite, this mean was 5.20 (Figure 1B-C). As for the length of the roots per cutting, the mean was 0.64 cm for peat and carbonized pine bark, against 5.83 cm for peat and expanded perlite (1:1).

Spring sprouting was obtained with the peat and expanded perlite substrate, reaching 33% (Figure 1C). The percentage of survival of the cuttings was 100% in both treatments.

It was also found in this study that substrates with density in the range of 70–110 g L^-1 (Table 2) were the ones that promoted greater root length. On the other hand, Vivatto Plus was the one that provided the highest percentage of rooted cuttings (90%), differing significantly from the others. The Vivatto Plus substrate used in this work has a density of 240 g L^-1, which is considered relatively low compared to the density of the sand, but it is more than twice the density of the peat and expanded perlite substrate (110 g L^-1) (Table 1).

3.2. Experiment 2 (E2)

Although the experiment was conducted in autumn, 90% of the cuttings cultivated in the Vivatto Plus substrate successfully took root. For the number of roots per cutting, there was no statistical difference between the substrates, obtaining a mean of 2.70 roots per cutting. However, there was a statistical difference in the mean root length. The mixture containing vermiculite and carbonized rice husk was superior, with a mean of 3.32 cm against 1.53 and 1.54 cm for sand and Vivatto Plus, respectively (Table 2 and Figures 1D-F).

During the autumn experiment, there was no sprouting on the tested substrates. The mortality percentage was 7% for the vermiculite and carbonized rice husk substrate and zero for the sand and Vivatto Plus substrate. As for the price of each substrate per cutting, Vivatto Plus was the most expensive of this experiment, costing R$0.15 per cutting (Supplementary table). Sand and the mixture of vermiculite and carbonized rice husk were the cheapest substrates among all evaluated in both seasons.

3.3. Anatomical analysis

The stem of S. setaceus presents a uniseriate epidermis and stomata with large substomatal chambers (Figure 2A-B). The cortex possesses a hypodermis formed by 4–5 layers of collenchyma located below the epidermis (Figure 2B-C) with a thick cuticle (evidenced by Sudan III) (Figure 2D). In the regions where the periderm is formed, no adventitious roots were observed (Figure 2E). In addition to starch (Figure 2F), the cortex is characterized by bulky parenchymal cells, with the presence of raphides, calcium oxalate druses and prismatic crystals (Figures 2G-I) and mucilaginous cavities (Figures 2A, J and K). In the cortex region, cortical bundles are also observed (Figures 2A and N). In the region of the vascular system, the vascular cambium forms a continuous ring, showing from the centre to the periphery: primary xylem, secondary xylem, vascular cambium (where fascicular and interfascicular cambium regions are recognizable), secondary phloem and perivascular fibres (Figure 2A, L-M). The adventitious roots of S. setaceus originate from the interfascicular cambium (Figure 3B). The adventitious roots cross the perivascular fibres (Figure 3B), the cortical parenchyma (Figure 3C), cortical bundles, secretory canals, the collenchyma, and the epidermis (Figures 3D-E).

Figure 2
Stem anatomy of pitaya-do-cerrado (Selenicereus setaceus) stem (cross sections): (A) General view of the stem, showing: pith (pi); secondary xylem (x2), vascular cambium (yellow dotted line) and secondary phloem (blue dotted line); perivascular fibres (fi); cortical bundles (red circles) and mucilaginous cavities (mu). (B) Epidermis (ep); stomata (st) and collenchymal hypodermis (co). (C) Collenchymal hypodermis (ruthenium red). (D) Cuticle (ct) (sudan III); epidermis (ep). (E) Periderm (pe). (F) Starch grains (lugol). (G) Raphides (black arrowhead). (H) Druse (dr). (I) Prismatic crystal. (J) Mucilage (white arrowhead). (K) Mucilaginous cavities (mu): cortex (ct) and Pith (pi). (L) Primary xylem (xy); secondary xylem (x2): ray cells (ra); vessel element (ve); fibres (fb); vascular cambium: fusiform (fu) and ray initials (ri). (M) Perivascular fibres (fi); secondary phloem (f2); vascular cambium (yellow dotted line) and secondary xylem (x2). (N) Cortical bundle: vessel (ve).

Figure 3
Morphoanatomical aspects of the adventitious roots of pitaya-do-cerrado (Selenicereus setaceus). (A) Newly formed roots in the stem; dotted white line emphasize the region where there is no apparent root (B/C) and the region with roots (detailed in D/E). (B) Developing root (ro) pushing the perivascular fibers (fi); radial system (rs) and axial system (as); mucilaginous cavities (mu); druse (dr). (C) Root in a more advanced stage of growth crossing through the cortex (ro). (D) Cross-section of the stem showing root that has just broken the epidermis (r1); root that has already broken the epidermis (r2); mucilaginous cavities (black arrowhead). (E) Root (ro) rupturing collenchyma (co) and epidermis; xylem detail (black arrowhead).

4. Discussion

We observed fewer and shorter roots in Selenicereus setaceus during autumn compared to spring when no sprouting of the cuttings occurred. According to Lone and Takahashi (2019), there is a decrease in rooting of Selenicereus undatus cuttings in the colder months. According to Neves et al. (2006), shorter days during autumn alter the physiological processes of matrix plants, such as photosynthesis and the transport of compounds, which hinder the rooting of cuttings. Thus, for the root development of the pitaya, higher temperatures favour the number of rooted cuttings, their development and consequent sprouting. Increased photoperiod also influences rooting, increasing root quality (branched shape and soft and flexible texture) as well as rooting percentage (Couvillon, 1988).

In the literature, studies indicate that substrate density is a factor that influences root length and should, therefore, be considered for cutting preparation (Peña-Baracaldo et al., 2018), even in easily rooted species such as S. setaceus. Denser substrates, such as sand, may pose higher resistance to root penetration due to compaction, which impairs the elongation capacity of the roots (Kämpf et al., 2006), as also observed for S. setaceus. In this species, the greatest root lengths were observed when using low-density substrates such as perlite and carbonized rice husk. These substrates, due to their higher porosity and lower susceptibility to compaction, provide more favourable conditions for root elongation (Kämpf et al., 2006).

When selecting a substrate, Fermino (2002) emphasizes that density is the most important physical property to consider. He notes that for smaller containers, lower substrate density should be used to prevent inhibition of root growth. For Selenicereus setaceus cuttings, we used multicellular trays and observed the best results with lower-density substrates (Tables 1 and 2).

Despite the excellent rooting capacity of the species in all substrates tested, sprouting was only observed in spring in the peat plus perlite substrate (Table 2). There are several reasons why some cuttings may develop roots but not sprout, as observed in most treatments in the present study (see Table 2). One of them is the need for hormones or even the deficit of nutrients in the substrate (Souza et al., 2024). According to Hartmann et al. (2017), the presence of nutrients in the substrates can influence both shoot development and the number and length of roots in cuttings.

Another important result in the present study concerns the low mortality of the cuttings of the species evaluated here. As expected, S. setaceus can survive different stress conditions for extended periods due to the presence of mucilage and starch, as evidenced here by the anatomical study (Figures 2 J-K). Mucilage and starch give cacti a great capacity to store water and nutrients in their stems, which are also photosynthetic (Gibson and Nobel, 1986). We emphasize that because it is a climbing cactus species of the presence of roots is an important mechanism to promote the anchoring of stems on the support (usually stems of other species), ensuring that the species reaches the canopy (Rowe et al. 2023; Soffiatti & Rowe, 2020). Thus, there is naturally an excellent rooting capacity in the species, which is primarily related to its fixation in the support, as described by Rowe et al. (2023) and Soffiatti & Rowe (2020).

As for the rooting anatomy, the adventitious roots that were formed in the cuttings originate in the interfascicular cambium – a novelty for the Cactaceae family. Some studies have linked the place of origin of root primordia to the rooting capacity of the species (Guan et al., 2020; Steffens and Rasmussen, 2016). In this context, in species that have easy rooting, the root primordia originate in the cambium region (Steffens and Rasmussen, 2016), as observed here for S. setaceus, although without specifying which cambium region. In Cactaceae, there is no record in the literature about the origin of adventitious roots. However, Haissig (1973) states that the adventitious roots in Cactaceae start more often from cells immediately adjacent to the primary xylem.

In regions where the periderm has already differentiated, adventitious root formation has not been observed. This suggests that the periderm, due to its thick, lignified cell walls, may act as a barrier to adventitious rooting. Therefore, for the production of Selenicereus setaceus seedlings via cuttings, it is recommended to select cuttings from regions of the matrix where the periderm has not yet replaced the epidermis.

In Selenicereus setaceus, it is recommended to produce cuttings during spring, as rooting occurs more rapidly compared to autumn, with sprouting also observed. In spring, all the tested variables performed better with the peat plus expanded perlite substrate. Pericyclic fibres and other tissues with thickened walls, such as collenchyma, do not hinder rooting in S. setaceus. However, the periderm may act as a barrier to rooting, so propagule collection from areas where the periderm is present should be avoided. Notably, adventitious roots form from the interfascicular cambium region, marking the first such report for this plant family.

Supplementary Material

Supplementary material accompanies this paper.

Supplementary table

Average price of the five substrates used in the December 2023 experiment (spring) and in the April 2024 experiment (autumn).

This material is available as part of the online article from https://doi.org/10.1590/1519-6984.293866

Acknowledgments

To Coordenação de Aperfeiçoamento de Pessoal de Nível Superior-Brasil (CAPES)-Finance Code 001 providing Doctoral Scholarship to first author. and to the Postgraduate Program PGAPV-UFPR.

Data Availability Statement

The entire data set that supports the results of this study was published in the article itself.

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Edited by

• Editor:
  Jairo Lizandro Schmitt

Publication Dates

• Publication in this collection
  17 Oct 2025
• Date of issue
  2025

History

• Received
  31 Jan 2025
• Accepted
  13 July 2025