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Anatomical characterization of pilosocereus pachycladus F. Ritter roots

Article  in  Acta horticulturae · June 2015

DOI: 10.17660/ActaHortic.2015.1087.29

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Anatomical Characterization of Pilosocereus pachycladus F. Ritter Roots
Alex S. Barbosa1, Rafaela D. Sá2, Flávia C. L. Silva3, Karina P. Randau2, Alberício P.
Andrade1 and Robson L. S. Medeiros1
1 2 3
Universidade Federal da Universidade Federal de Universidade Federal
Paraíba Pernambuco Rural de Pernambuco
Programa de Pós- Programa de Pós-Graduação Departamento de
Graduação em Agronomia em Inovação Terapêutica Biologia
Areia, Brazil Recife, Brazil Recife, Brazil

Keywords: anatomy, cactus, Facheiro

Abstract
Pilosocereus pachycladus F. Ritter is a native cactus which is grown in the semi-
arid area of Northeastern Brazil. The specimens have an ornamental value, yet studies
on their anatomy are scarce. The root anatomy of P. pachycladus was studied in order
to understand better the adaptation mechanisms. Specimens were collected in Areial,
Arara and Boa Vista, State of Paraíba. Cross sections of roots were clarified with
sodium hypochlorite (50%), stained with safranin and blue astra and examined under
light microscopy. In general, the secondary structure of the roots was similar for the
genotypes collected at the three sites. We observed the replacement of the epidermis
by periderm, a cortex with few layers of parenchyma, collateral vascular bundles and
fibers located in the parenchyma rays between beams. Druses were present in the
cortical region. These were observed in higher numbers in roots collected in Areial
and less frequently in roots from plants growing in Boa Vista. The presence of crystals
indicate the adaptive capacity towards the xeric conditions, as a higher amount of
druse was observed in specimens growing in warmer climate. This study contributes
to the establishment of anatomical parameters, contributing to the taxonomy of the
species.

INTRODUCTION
The Cactus family occurred throughout the tropical America region with
approximately 1,900 species, distributed in around 100 genera (Areces, 2004). Only
Rhipsalis baccifera species occurs naturally in tropical Africa and Sri Lanka. The family is
divided into three subfamilies: Pereskioideae, Opuntioideae and Cactoideae (Nyffeler,
2002). Cacti have a broad adaptation to arid and continental semiarid environments.
Barthlott and Hunt (1993) describe them as evergreen trees, shrubs or vines, with juicy
stems and adaptations both morphological and in the floral structure for different types of
pollination.
Brazil is considered as the third largest center of diversity of Cactaceae, with about
200 species (Souza and Calvente, 2013). The savanna is identified as diversity center of the
Cereeae tribe, while the north central part of Mexico and the semiarid region of the western
United States is characterized by the presence of species from the Cactae and Pachycereeae
tribes. In South America, the Andean region, the representatives of Browningieae,
Notocacteae and Trichocereeae (Barthlott and Hunt, 1993) characterize an area occupied
by Peru, Bolivia, Chile and Argentina.
The facheiro belongs to Pilosocereus genus and is found in Mexico and South
America. The Pilosocereus genus (Byles & Rowley), comprises 36 neotropical species
belonging to the subfamily Catoideae and Cereeae tribe (Zappi, 1994). In Brazil,
approximately 26 species are found, mainly distributed in the caatinga, on rocks and dunes,
in rocky outcrops, in the thorn and wild forests and in grasslands in Brazilian Highlands
(Zappi, 1994). According to Zappi and Taylor (2008) in total 160 cactus species are
growing in Brazil, 42 species, representing 26% of the family occur in rocky fields, while
31% are distributed in the savanna.
The native cacti, especially the Pilosocereus genus, is an important regional plant
genetic resource in order to maintain the ecological niches and the remaining savanna
fragments. Therefore, studies on the dispersion patterns, they relations with the
environment and their adaptive capacity to the semiarid environment are relevant. Cacti has
an important value as an ornamental plant, especially because they are distinctive plants in
terms of growth and morphological characteristics. The native cacti from Brazil still need
more in depth studies aiming to understand relations between plant and environment.
Anatomical, morphological and physiological characteristics of species of Cactaceae show
adaptations to arid and semi-arid environments and mechanisms that lead these plants to
interact with their environment (Gibson and Nobel, 1986).
Among the most significant stem anatomical changes, Terrazas and Mauseth (2002)
highlighted the presence of epicuticular wax, thick cuticle, stomata located in the epidermis
depressions (in some species), collenchymatic hypodermis and the development of large
proportions of parenchymal tissue with mucilage cells. However, studies of comparative
anatomy are needed to clarify and gather information for the characterization of
Pilosocereus pachycladus F. Ritter. In this study we described the anatomy of roots of P.
pachycladus coming from the caatinga Paraíba, Brazil.

MATERIALS AND METHODS

Plant material
The plant material was collected in Arara (25 M 192007 9243179 UTM), Areial (25
M 175834 9218837 UTM) and Boa Vista (24 M 698981 9133272 UTM), State of Paraíba,
Brazil. All material was analyzed and identified as Pilosocereus pachycladus F. Ritter
(Cactaceae) and was deposited in the Herbarium Jaime Coelho de Moraes - EAN (Number:
20101, 20102 and 20103, respectively).

Microscopic analysis
Cuts were made in cross sections in the midline of fresh roots, freehand, using
common razor blade and petiole marrow timber (Cecropia sp.) as support material was
used. The sections were clarified with 50% sodium hypochlorite solution and stained with
safranin and astra blue (Johansen, 1940). Later, semi-permanent histological slides were
prepared, containing cross-sections of the plant material, following the usual procedures in
plant anatomy (Johansen, 1940; Sass, 1951; Silva et al., 2013). Digital images were
captured by optical microscope (Alltion®) coupled with digital camera by Toup program
View Image.

RESULTS AND DISCUSSION

The roots internal structure in the secondary stage were similar in all the individuals
analyzed (Fig. 1A-1C). The coating system of Pilosocereus pachycladus roots is
constituted by the periderm (Fig. 2A), which completely replaces the epidermis. The
periderm cells have rectangular shape and the cell walls are strongly thickened. The cortical
region is reduced, due to the development of the vascular system and periderm (Fig. 2A).
A similar internal constitution of the roots was also observed in P. pachycladus subsp.
pernambucoensis (Ritter) (Zappi, 1999) and five other species of Pilosocereus (Silva and
Alves, 1999), as well as in other cactus species i.e. Melocactus × horridus Wedermann
Notizbl (Arruda et al., 2005).
The vascular bundles are collateral, ray parenchyma were observed separating the
vascular bundles, which, in cross section, have 3 to 4 rows of cells (Fig. 1A-1C). At some
points, the xylem forked and new rays were formed (Fig. 1A and 1B). According to Silva
and Alves (1999), in P. tuberculatus (Werderm.) Byles & Rowley, the number of rays of
cell rows can reach six. According to these authors, the parenchyma rays increase the
amount of specialized cells for storing water. By this it is an important characteristic of
savanna species for adaption to the limited availability of water.
Fiber isolated groups displaying as mentioned were observed (Fig. 2B). The
presence of amyloplasts also have been observed in the Cactaceae family (Silva and Alves,
1999; Arruda et al., 2005). However, so far they were not displayed in this specie.
The abundant occurrence of calcium oxalate grouped crystals in Cactaceae is reported
by Metcalfe and Chalk (1972). In P. pachycladus, druses were found in parenchyma rays
(Fig. 2C) and in the cortical region (Fig. 2D). However, they were observed in higher
numbers in roots collected in Areial compared to roots of specimens collected in Boa Vista.
According to the literature, the presence of crystals might indicate the adaptation capacity
to the xeric conditions (Fahn and Cutler, 1992). This is in agreement with our findings in
this approach, where we observed a larger amount of druses in the genotypes growing in
warmer climate conditions.

CONCLUSION
P. pachycladus has adaptations to the environment as druses and a thick layer in the
epidermis. These adaptations serve to survive the long periods without water during the dry
season in the caatinga. This approach contribute to the establishment of anatomical
parameters and to the taxonomy of the species.

Literature cited
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 Figures

Fig. 1. Roots in secondary structure Pilosocereus pachycladus F. Ritter: Overview of

Pilosocereus pachycladus roots cross-sectional collected in: A) Arara; B) Areial; C) Boa
Vista, PB - Brazil. Bars: A, B, C = 100 µm.

Fig. 2. Cross sections of Pilosocereus pachycladus F. Ritter roots: A) Cortical region and
periderm; B) fibers between parenchyma rays; C) Druze in parenchymatous radius; D)
Druze in the cortical region. Bars: A, B, C, D = 500 µm.

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