Acta Botanica Brasilica

Print version ISSN 0102-3306

ActaBot.Bras.vol.18no.4SoPauloOct./Dec.2004
http://dx.doi.org/10.1590/S0102-33062004000400004

Tracheary elements of five taxa of Cactaceae of

Pernambuco savanna, Brazil

Traqueary elements of five taxa of Cactaceae of the calabash

from Pernambuco State, Brazil

Emilia Cristina Pereira de Arruda

Melo-de-Pinna III

I, *

; Marccus Alves

II

; Flavia Gladys

Master's student at the University of So Paulo II Department of Botany, Institute

of Biology, Federal University of Pernambuco, Av. Moraes Rego s / n, Ciudad
Universitaria, CEP 50632-970, Recife, PE, Brazil III Institute of Biosciences,
University of So Paulo, C. Postal 4005, CEP 0161-970, So Paulo, SP, Brazil
I

ABSTRACT
This paper presents the morphological characterization of vessel elements and
WBT ("tracheid with wide thickening") root and cladodes five taxa of Cactaceae
( Harrisia adscendens (Gurke) Britton & Rose, melocactus horridus Wedermann
Notizbl., M . zehntneri ( Britton & Rose) Luetzelb., Tacinga inamoena (Schumann)
NP Taylor & Stuppy and T . palmadora (Britton & Rose) NP Taylor & Stuppy)
occurring in a stretch of caatinga of Pernambuco State, Brazil. The vessel
elements in roots as much in the cladodes have practically identical among the
taxa studied, pits alternate, opposite and scalariform being observed, and simple
transverse and oblique perforation plates. The tracheids (WBT) were recorded
only in the cladodes four taxa studied, except Harrisia adscendens . The WBTs
have annular or spiral thickening and thickening occurs only in
mixed melocactus . The tracheal elements were measured, showing that
in melocactus both the root and the cladodes, feature elements of long vessel and
the WBT as these are higher in cladodes of melocactus .
Keywords: anatomy, Cactaceae, calabash, tracheal elements

ABSTRACT
This work brings the morphological characterization of traqueary elements
(vessels and tracheids) in root and cladode of five taxa of Cactaceae ( Harrisia
adscendens (Gurke) Britton & Rose, melocactus horridusWedermann
Notizbl., M . zehntneri (Britton & Rose) Luetzelb. , Tacinga inamoena (Schumann)
NP Taylor & Stuppy and T . palmadora (Britton & Rose) NP Taylor & Stuppy)
occurent in the caatinga of Pernambuco State, Brazil.The vessels Observed
elements in the roots and cladodes seem almost the same among Studied taxa,
were Observed alternate, opposite and scalariform pitting, and simple perforation
plate oblique or transverse. The tracheids (WBT) were exclusively Noted in the
cladodes of four Studied taxa, except adscendens Harrisia . The WBTs with
annular and helical thickening are present in all taxa and the mix thickening is
present only inmelocactus . The vessel elements and tracheids were showing
Measured and que the vessel elements are longer in roots and cladodes
of melocactus and the longest WBTs were Observed only in cladode
of melocactus .
Key words: anatomy, Cactaceae, calabash, traqueary elements

Introduction
The vascular system of the Cactaceae is basically constituted by conducting cells
of the xylem and phloem as in other groups of vascular plants. But in Cactaceae
usually the xylem tissue have the highest proportion of parenchyma cells
compared to cells lignified walls (vessel elements and fibers) thus composing the
aquifer called parenchyma (Soffiatti & Angyalossy 2003; Mazzoni-Nurseries &
Costa 2003).
Some of the main characters of the vessel elements of Cactaceae include pits
scalariform, alternating the presence of irregular and simple perforation plates
(Metcalfe & Chalk 1950, Gibson 1973, 1977; Gibson & Nobel 1986; Soffiatti &
Angyalossy 2003).
In addition to vessel elements, fibers and parenchyma cells, some species of
Cactaceae may present in the xylem, a special type of tracheid called "wide-band
tracheid" - WBTs (Metcalfe & Chalk 1950; Gibson 1973; Mauseth 1999). This cell
type is characterized by the absence of perforations in their walls, bands of rigid
secondary thickening, which may be annular, helical or double helix, directed to
the heat cell (Gibson, 1977; Mauseth 1989; Landrum 2001).
There are still few studies conducted with WBT, so that its origin and function are
still uncertain (Conde 1975), although some researchers relate their occurrence
with juiciness (Boke 1944; Mauseth 1993). However, few studies have been
developed to demonstrate that these cells may be important for the phylogenetic
relationships of the Cactaceae (Mauseth et al 1995;. Mauseth & Landrum 1997;
Landrum 2002). The dimensions of vessel elements and fibers, some work has
been done (Mauseth 1993; Mauseth & Plemons-Rodriguez 1998). Nobel & Gibson

(1986) reported a correlation between length and diameter of these cells with
habit of plants.
The aim of this work is to characterize the tracheal elements (vessel elements
and WBT) five taxa of Cactaceae occurring in a stretch of caatinga of Pernambuco
State, and through these observations, seek these cells detect diagnostic
differences between the organs (roots and cladodes) and between the taxa
studied.

Material and methods

This work was developed in the city of Alagoinha (08 29'' S and 36 47'' W),
about 780m.sm (FIAM 1994), Cashew Farms in Dry and Lagoa Seca, located on a
stretch of caatinga of Pernambuco, in boundary between the Central and the
Wasteland Hinterland as proposed by Andrade-Lima (1957, 1960)
classification. Excursions were made to the study area for collection of vegetative
organs (roots and cladodes) of selected taxa and reproductive well as field
observations.
The selection of taxa was done through consultation with the collections available
in local herbaria (Herbarium of the Federal University of Pernambuco - UFP;
Herbarium of the Federal Rural University of Pernambuco - PEUFR and Herbarium
Pernambucana Agricultural Research Corporation - IPA). Due to the abundance in
the study area, variety of ways and in different subfamilies these species were
chosen: adscendens Harrisia (Gurke) Britton & Rose, with procumbent
habit; melocactus horridus . Wedermann Notizbl, M . zehntneri (Britton &
Rose) Luetzelb, both globular habit;. Tacinga inamoena (Schumann) NP Taylor &
Stuppy and T . palmadora . (Britton & Rose) NP Taylor & Stuppy both with a
shrub H. adscendens , melocactus horridus and M. zenhtneri are
representatives of the subfamily Cactoideae while inamoena
Tacinga and T. palmadora subfamily Opuntioideae (Hunt & Taylor 1990). The
classification of the habit of the taxa studied follows the proposal by Gibson &
Nobel (1986) for the Cactaceae family.
After collection of the material was intended for the manufacture of dried
specimens, according to the usual rules (Mori et al. 1989) and deposited in the
Herbarium UFP. Part of plant material (roots and cladodes) was collected and
fixed in FAA 50, after 24 hours, transferred to 70% ethanol (Johansen 1940), for
its preservation. From this material, the technique of dissociation, both roots of
cladodes, with a solution of hydrogen peroxide and acetic acid 1:1 (Franklin
1945) was used. After the material loose natural color, it was washed with
distilled water until complete removal of the dissociation solution and stored in
70% ethanol (Franklin 1945). Fragments of this material were macerated on
histological slide, with added drops of 1% safranin in 50% ethanol (Kraus &
Arduin 1997) for staining and mounted in 50% glycerin (Purvis et al . 1964).
The analysis of vessel elements and WBT was performed under a light
microscope. The measurement was made in the light microscope coupled
millimeter eyepiece. For registration of the cells, the slides were photographed in
the light microscope and schematic representations were made on camera lucida
attached to a light microscope.
Statistical analysis was performed with BioEstat 2.0 software (Ayres et al. 2000).

Results
The vessel elements of the taxa analyzed, both in roots and in cladodes, are
characterized by having simple perforation plates with transverse or oblique,
scalariform pits, alternate and opposite and also pontoados appendages ( Fig. 1AC , 2A- U ). In Figures 3 and 4 it is possible to observe the variation in length of
these cells in roots and cladodes, respectively.

Statistical tests showed that longer vessel elements observed in the roots belong
to melocactus , and this difference was statistically significant (p <0.05) in almost
all combinations performed, except between H.adscendens and T. inamoena ( Fig.
3 ). In cladodes, the highest values found for the vessel elements occur
in H.adscendens and melocactus , in particular M. zehntneri , which is also
statistically significant relationship in almost all combinations performed, except
between M. horridus and M. zehntneri ( Fig. 4 ).
Only in the xylem of cladodes of melocactus horridus , M. zehntneri , inamoena
Tacinga and T. palmadora , which are among the most succulent taxa studied,
WBT were observed, in adscendens Harrisia , the cladodes is less juicy. The rigid
bands of secondary thickening present in these cells are shown of three types:
spiral ( Fig. 1D ), annular ( Fig. 1D ) and mixed ( Fig. 1F ), the annular
thickenings and helical occur in the same cell. The WBT with spiral and annular
thickening were observed in melocactus horridus , M. zehntneri , inamoena
Tacinga and T. palmadora , while mixed with thickening were registered only in
the cladodes of melocactus . You can also notice variations on the length of WBT
among the taxa studied ( Fig. 5 ), and the longer occur inmelocactus , especially
in M. horridus . This relationship is statistically significant for all combinations
performed.

Discussion
The vessel elements of cladodes and roots of the taxa analyzed are presented
essentially as described in the literature of Cactaceae (Metcalfe & Chalk 1950;
Gibson 1977; Soffiatti & Angyalossy 2003), with no specificity types among the
taxa studied.
As for dimensional patterns, it was observed that the length of the vessel
elements of the taxa analyzed in this study is within the range expected for
Cactaceae (100-500m) as mentioned by Metcalfe & Chalk (1950) values.
Gibson & Nobel (1986), through analysis of 119 species of 50 genera of the
subfamily Cactoideae and 35 species from 17 groups Platyopuntias, found the
correlation between the lengths of vessel elements and fibers with plant size, and
these Longer the greater the size of the plant cells. However, Carlquist (1975)
and Zimmerman (1983) showed the occurrence of vessel elements shorter shrub
species in arid regions, such as is found in most of the Cactaceae. In taxa studied
there seems to be a correlation between the length of vessel elements and plant
habit, and in view of a tendency of these cells were longer in both roots and in
cladodes in melocactus , smaller plants analyzed. Although H. adscendens plant
larger study, using arguments longer vase in the cladodes, it is possible to
observe similar pattern in M. zehntneri .
Earl (1975) showed length of elements of relatively constant vessel in five species
of Opuntia analyzed by him.According to the author, this may be due to the
relative similarity between species. This may in part be corroborated in the
present study compared the two taxa Tacinga ( Tacinga inamoena -132m and T.
palmadora- 161m) and the two of melocactus ( M. horridus - 185m and M.
zehntneri - 198m), although the number of representatives of each gender is
small.
Bailey (1957) mentioned general evolutionary trend of reducing the length of
vessel elements and fibers in Cactaceae. Based on this character plants of this
study, which would be derived from the Tacinga are actually derived as less
than melocactus (Hunt & Taylor 1990) Elements which have longer
vessel. According Mauseth (1993), short glass elements prevent embolism and
the consequent interruption of driving, especially on plants of dry environments,
such as those studied here.

The occurrence of WBT in Cactaceae is mentioned by several authors (Metcalfe &

Chalk 1950; Gibson & Nobel 1986; Mauseth 1999; Landrum 2002). The origin
and function of these cells are still uncertain (Conde 1975), although some
authors correlate their occurrence with the degree of succulence of the plant
(Boke 1944; Gibson 1977; Fahn & Cutler 1992; Mauseth 1993), while others with
the same evolution (Mauseth et al. 1995; Landrum 2002). According to Gibson
(1977), when these cells are present so there is a tendency to reduce the
proportion of vessel elements, and to the absence of xylem fibers. Although WBT
have allowed the distinction of taxa analyzed, bearing in mind its absence only
in H. adscendens , the claims made by the authors cited above suggest that
studies should be conducted in these and other taxa to prove the real function of
these structures.
Due to WBT occur in excessively succulent tissues, possible because of the strict
bands of secondary thickening is strengthening this tissue, aiding in support
(Bailey, 1966, cited in Conde 1975). According Mauseth et al.(1995), these bands
prevent the collapse of the primary cell walls keeping fit to drive even in periods
of water stress fire. Also according to the author, because of these properties,
WBT may represent important adaptations to succulent plants in xeric
environments. The thickening occurring in these cells may be annular, helical or
double-stranded (Metcalfe & Chalk 1950; Gibson & Nobel 1986; Landrum
2001). Gibson (1977) mentioned the occurrence of WBT typically narrow and
slender and with spiral thickening Cactoideae, although the annular thickening
can occur in small representatives of that subfamily. Have the Opuntioideae,
although presenting narrow and fusiform WBT, the thickening of these cells in
these species is annular or, rarely, ring-worm. There is no specificity between
types of thickening of WBT among the taxa studied, since all have annular and
helical thickening, except for the ring-worm, described here as mixed, which
occurs only in melocactus .
Based on the results, we suggest that the tracheal elements observed here
corroborate the results for forms and types described for the Cactaceae. Due to
the similarity of these elements in roots and cladodes, as well as among the taxa
studied, it is not possible to diagnose the same based on this character. As the
WBT are absent in the roots of all taxa studied and cladodes of H. adscendens ,
these may represent a diagnosis or even be related to the juiciness
character. Regarding the pattern of thickening, restricted to cladodes mixed
typemelocactus also allows the diagnosis of the taxa studied.
However, it should be emphasized that the small number of taxa analyzed,
necessitates the expansion of studies on the tracheal elements in other family
representatives, aiming to fix the characters observed in these taxa, and establish
other also useful to diagnosis.

Thanks
The authors thank Dr. Nigel Taylor, for his kindness in identifying botanical
material; PIBIC / CNPq / UFPe (Proc. 20310), the granting and renewal of
scientific initiation scholarship granted to the first author.

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