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Morphology and Anatomy of the Flowers of
Trichilia catigua A. Juss., T. elegans A. Juss. and
T. pallida Sw. (Meliaceae)
Article in Brazilian Archives of Biology and Technology · December 2001
DOI: 10.1590/S1516-89132001000400008 · Source: DOAJ
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383
Vol. 44, N. 4 : pp. 383 - 394, December, 2001
ISSN 1516-8913
BRAZILIAN ARCHIVES OF
BIOLOGY AND TECHNOLOGY
Printed in Brazil
A N
I N T E R N A T I O N A L
J O U R N A L
Morphology and Anatomy of the Flowers of Trichilia catigua
A. Juss., T. elegans A. Juss. and T. pallida Sw. (Meliaceae)
Luiz A. de Souza*; Ismar S. Moscheta; Káthia S. M. Mourão and Adriano Silvério
Universidade Estadual de Maringá, Departamento de Biologia, Avenida Colombo, 5790, 87020-900 Maringá - PR,
Brazil
ABSTRACT
Morphological and structural features of flowers of Trichilia catigua A. Juss., T. elegans A. Juss. and T. pallida Sw.
(Meliaceae) are presented here with the purpose to stimulate future reproductive and preservation studies of these
brazilian native species. Anthesis occurred from April to August in T. catigua, January to May in T. pallida and
September to January in T. elegans. Sepals and petals presented a papillose and pilose epidermis and a
parenchymatous mesophyll. Male flowers presented pistillodes with abortive ovules and a staminal tube with
tetrasporangiate anthers. Anther wall had a papilose epidermis, a fibrous endothecium, two middle layers and a
secretory tapetum. Female flowers presented antherodes and bi - or tricarpellate syncarpous pistils. Ovary had a
simple structure with anatropous, bitegmic ovules. Style was hollow or solid and the stigma was constituted by uni or bicelullar trichomes. Strictly dioecious specimens were not found in the study area.
Key words: Trichilia, Meliaceae, floral morphology and anatomy, flowering
INTRODUCTION
Trichilia P. Browne, belonging to the Meliaceae
family, accounts for about 70 species distributed
throughout Tropical America. The species
frequently present unisexual (bisexual to a lesser
degree) flowers in dioecious plants and there are
even records of polygamous plants (Pennington,
1981; Klein, 1984). In the forest remnants in and
near Maringá, Paraná State, Brazil, three species
of Trichilia occur: T. catigua A. Juss (catiguá), T.
elegans A. Juss. (pau-de-ervilha) and T. pallida
Sw. (baga-de-morcego). These species have a
wide distribution in South and Central America.
The first two are more abundant in southern Brazil
and the last one is dispersed throughout Brazil and
be very common in the southeastern gallery forests
(Pennington, 1981; Klein, 1984).
*
The flowers of Trichilia species are not well
known in their anatomical aspects. However,
anatomical features such as ovaries and ovules in
T. grandifolia Oliv., T. elegans A. Juss., T. pallida
Sw. and T. catigua A. Juss. are described by
Boesewinkel (1981), Beltrati (1984), Beltrati and
Brunini (1988) and Moscheta (1995), respectively.
Morphological descriptions of flowers are limited
to the works of Pennington (1981) and Puentes et
al. (1993). Studies about floral biology include
works of Styles (1972), Opler and Bawa (1978)
and Opler (1983).
The process of the human occupation of Paraná
decimated the original forests of the state. Forest
coverture has reduced from 83.41% to
approximately just 7% presently, most of which is
concentrated in the “Serra do Mar” and Iguaçu
National Park. In the northwest region of Paraná,
Author for correspondence
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384
Souza, Luiz A. de et al.
the situation is more critical, with less than 1% of
the forest remnants located in units of conservation
and on the islands in the Upper Paraná River
which are subject to flooding (Ipardes, 1992;
Campos, 1997).
With the purpose of understanding the
differentiated behavior of Trichilia plants
(dioecious, monoecious and polygamous) recorded
in the literature and stimulating future
reproductive and preservation studies on Paraná’s
forests, morphological and anatomical features of
the flowers of Trichilia catigua A. Juss., T.
elegans A. Juss., and T. pallida Sw. are presented
and discussed.
RESULTS
Morphology
The flowering time for the species can be long, as
in T. catigua, where the floral buds can stay on the
plant for five to six months before the anthesis
process begins. In T. pallida and T. elegans the
floral buds stay on the plants for about two
months. The anthesis occurred from April to
August in T. catigua, from January to May in T.
pallida and from September to January in T.
elegans.
The flowers of the species were minute,
approximately three millimeters long in T.
elegans, about four millimeters in T. catigua
and a little more than five millimeters in T.
MATERIALS AND METHODS
pallida. These flowers occurred in axillary
panicles in the three species. However, the
The “Horto Florestal Dr. Luiz TeIxeira Mendes”,
inflorescence in T. catigua was congested and
where the species were observed and studied, is
in T. pallida small racemes with short axes
situated in the city of Maringá, Paraná State,
can
occur. The flowers presented a short
Brazil between the meridians 51° 30' and 54°W
and the parallels 22° 30' and 24° 30' S, with an pedicel, at whose base occurred a reduced
bract and two bracteoles with the same shape,
average altitude of 556 meters. In this forest
remnant, there are shrubs and small trees of
but smaller dimensions. They were still
Trichilia elegans A. Juss., and small trees of T.
dichlamydeous, actinomorphic and unisexual
catigua A. Juss. and T. pallida Sw., whose
(Figures 1 and 2).
vouchers were deposited in the Herbarium of the
Universidade Estadual de Maringá, under numbers
HUM 5111, HUM 2210 and HUM 197,
respectively.
The flowering time for the species was followed in
at least five previously marked specimens between
April 1999 and June 2000 by means of weekly
observations. During this period, flowers were
collected before and after the anthesis,
immediately analyzed in the laboratory and fixed
in FAA 50 and FPA 50 (Johansen, 1940). The
anatomical study of the collected botanical
material was sectioned by microtome techniques
in paraffin wax (Johansen, 1940) or methacrylate
(Ruetze and Schmitt, 1986). Erlich hematoxylin
and safranin were used for staining (Johansen,
1940). In addition, temporary and semi-permanent
slides obtained from hand sections and stained
with safranin and astra blue were made. The
following stains were used for specific color tests:
phloroglucinol and sulfuric acid, Sudan IV and IKI
for lignins, lipids and starch, respectively
(Johansen, 1940; Berlyn and Miksche, 1976).
The synsepalous green calyx had four sepals in T.
catigua and T. pallida (Figure 21) and five in T.
elegans. In T. catigua the sepals were more pilose
and reduced. The corolla presented four separate
petals (Figure 21), except in T. elegans, which had
five separate petals, white to yellow in color and
lanceolate (Figures 3 to 5) to ovate in shape. The
petals of T. elegans presented only one central
vein, which branched out into secondary veins of
smaller caliber (Figure 4). In T. catigua the petals
showed two main veins, which emited several
branches (Figure 3). In T. pallida, several veins
started from the petal base. The central vein
presented larger caliber. Eventually, these veins
branched out (Figures 5 and 7).
In male flowers the androecium was constituted by
a staminal tube, which had tetrasporangiate anthers
(Figure 2). The tube apex of T. elegans had ten
anthers alternating with thin and pilose extensions.
There were seven or eight alternating anthers with
edges in T. catigua. These seven or eight pilose
anthers were attached by small filaments to the
tube. All the anthers had longitudinal opening. The
androecium presented the same morphology in
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Morphology and Anatomy of the Flowers
female flowers, but the anthers lacked pollen
(Figures 1 and 20).
The gynoecium in female flowers presented a
syncarpous pistil. The conical ovary (Figure 1) had
three carpels and locules, two ovules per locule
with axile placentation, short style and prominent
385
stigma. Some variations were also found in the
studied species: in T. elegans the gynoecium might
have present two or three carpels and there were
many shaggy trichomes in the ovary of T. pallida.
In male flowers, the gynoecium was represented
by a pistillode with abortive ovules (Figure 2).
Figure 1–5. 1–2 - Female flower of T. elegans and male floral bud of T. pallida in longitudinal section, respectively;
3–5: Cleared petals of T. catigua, T. elegans and T. pallida, respectively. (ad=antherode; an=anther; ne=nectary;
nv=vein; ov=ovary; pd=pistillode; pe=petal; se=sepal; sg=stigma; st=staminal tube; sy=style; vb=vascular bundle)
Nectaries were present only in T. elegans and T.
pallida (Figure 2). The nectaries in T. elegans, not
always present, occurred as a disk-like structure
involving the base of the ovary. In T. pallida the
nectaries were formed at the base of the ovary and
the staminal tube.
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Souza, Luiz A. de et al.
Anatomy
The sepals of the species had a single, papillose
epidermis (Figure 6), with unicellular trichomes of
thin extremity in T. catigua and T. pallida. The
parenchyma cells of the mesophyll showed thin
walls and vary in size and shape. There were also
secretory cavities and wide secretory cells (Figure
6). Vascularization was made by a central
collateral vascular strand, next to others with small
dimensions.
The petals had a uniseriate papillose epidermis,
with trichomes, stomata (Figures 7 to 13) and
cuticle, which could also be ornamented (Figure
11). The papils occurred frequently in the
epidermis of the three species and showed thick
cellular walls (Figures 8 to 11). The trichomes
could be uni or pluricelullar, with thin or thick
cellular walls and pointed, or more or less rounded
extremities (Figures 10 and 12). They were
common in the petal edges. In the floral buds there
was a perfect adjustment among the trichome
epidermes with contiguous petals (Figure 13). The
mesophyll was parenchymatic and constituted by
cells with thin walls and by secretory cells and
cavities. This region had few layers in T. catigua
and T. elegans (Figure 9) and many in T. pallida
(Figure 8). The presence of calcium oxalate druses
was notable in the parenchyma cells of T. catigua
and T. elegans. There was a central collateral
vascular strand and others of small dimensions
immersed in the mesophyll (Figure 7).The
nectaries were constituted by a single epidermis
with thin-walled cells and non-glandular
unicellular trichomes with relatively thick walls
(Figure 15), and by a secretory parenchyma where
the phloematic vascular tissue and secretory cells
and cavities stood out (Figure 14).
The male flowers of T. catigua and T. elegans
presented androecium constituted by a staminal
tube with a unilayered epidermis and prolonged
non-glandular
unicellular
trichomes.
The
epidermal cells were papillose and showed a
relatively thick and ornamented outer periclinal
wall. Below the epidermis there was a
homogeneous parenchyma with thick-walled cells
(Figure16). It presented secretory cells and
cavities and vascular bundles; one for each anther
(Figure 21). The vascular bundles of the tube
could be collateral or concentric. The androecium
of T. pallida also had a staminal tube, but differed
from the other two species by nectaries and papils
at the base and trichomes in the proximities of the
anthers; however, in T. pallida the papils did not
show a thick or ornamented wall. The wall of the
immature anther presented a single papillose
epidermis, endothecium with thin-walled cells,
slightly prismatic in shape, two middle layers and
a secretory tapetum with binucleate cells (Figure
18). In the mature anther, the endothecium cells
acquired thickenings on the anticlinal walls and on
the inner periclinal wall. The middle layers and the
tapetum began a disintegration process, were
flattened and crushed (Figure 19). The pollen
grains remained united in tetrahedron-type tetrads
(Figure 17).
In female flowers, the antherodes without pollen
had a papillose epidermis at maturity and an
endothecium with parietal thickenings. The middle
layers and the tapetum disintegrated in this phase
(Figure 20). The only pistil in female flowers had
an ovary with a uniseriate outer epidermis, which
showed stomata and a cuticle. The epidermal cells
varied from tabulate to prismatic in shape. The
epidermis was glabrate and papillose in T. elegans
and pilose in T. catigua and T. pallida. The
trichomes, very abundant in these species, were
non-glandular, unicellular, presented thick cellular
walls and thin extremities. In the multilayered
parenchyma mesophyll there were secretory cells
and cavities, notably numerous in T. elegans. A
single and glabrate epidermis covered the ovarian
cavity and presented cells tabulate to slightly
prismatic in shape. Both the subepidermal layers
(Figure 30) as the ventral epidermis of T. elegans
and T. pallida could undergo periclinal divisions
in the preanthesis stage. Ovarian vascularization
was made by differentiated vascular bundles or by
lateral, ventral or marginal and dorsal procambial
bundles, in accordance with carpel number. The
septs, which separated the locules, were
interrupted in the ovary apex and joined the
locules to each other (Figures 21 and 22).
The pistil still had a short style with an epidermis
similar to that of the ovary, which covered the
parenchyma tissue, where there were vascular
bundles and central transmitting tissue (Figures 23
to 25).
The vascular bundles varied in number, three
being of larger dimension (one for each carpel)
and other smaller ones might eventually be present
(Figures 23 and 24). In T. elegans specimens with
two carpels, there were only two vascular bundles
(Figure 25). The transmitting tissue was formed by
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Morphology and Anatomy of the Flowers
the epidermis and the secretory subepidermal layer
which were arranged in the central region of the
style as three small edges (Figures 23, 24 and 29)
or two in T. elegans specimens (Figures 25 and
28).
387
The style in T. catigua and T. pallida, represented
by a small rift, was hollow (Figures 23 and 29). In
T. elegans, however, a central rift occured in the
middle region of the style only, therefore
presenting a solid nature in the base and in the
apex (Figures 25 and 28).
Figure 6–11. 6 - Detail of the floral bud sepal of T. elegans in cross-section; 7-8: Diagram and detail of the floral
bud petal of T. pallida in cross-section; 9: Detail of the petal of T. elegans in cross-section; 10-11: Detail of the
adaxial and abaxial surface of the petal epidermis of T. pallida and T. elegans, in frontal view, respectively,.
(eb=abaxial epidermis; ed=adaxial epidermis; pa=epidermal papillae; sc=secretory cell; vb=vascular bundle)
The transmitting tissue traversed the whole style
and reached the apical region of the ovarian locule.
The stigma presented unicellular trichomes, more
frequent, and bicelullar; these trichomes came
from the immature epidermis of the stigma, whose
cells could undergo periclinal divisions and
posterior lengthening (Figures 26 and 27).
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Souza, Luiz A. de et al.
Figure 12-15. T. pallida. 12: Detail of the abaxial surface of the petal epidermis, in longitudinal section, showing
uni and multicellular trichomes; 13: Detail of the contiguous epidermis of the petals, showing the papillaes; 14:
Structural details of the nectary; 15: Nectary epidermis in frontal view. (bt=trichome base; sc=secretory cell)
Brazilian Archives of Biology and Technology
Morphology and Anatomy of the Flowers
389
Figure 16-20. 16 - Structural detail of the staminal tube wall of T.catigua; 17: Detail of the secretory tapetum and
the microspores in tetrads in T. pallida. 18-19: Detail of the young and mature anther wall in T. pallida and T.
elegans, respectively; 20: Diagram of the antherode in T. elegans, in cross-section. (de=dorsal epidermis; ve=ventral
epidermis; ep=epidermis; ie=immature endothecium; me=mature endothecium; ml=middle layers; as=pollen sac;
ta=tapetum; vb=vascular bundle)
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Souza, Luiz A. de et al.
Figure 21-25 - Diagrams of the cross-sections. 21: Flower of the T. pallida ; 22: Upper portion of the ovary in T.
catigua. 23-24: Basal and apical portion of the hollow style in T. catigua; 25: Solid style in T. elegans (de=dorsal
epidermis; ve=ventral epidermis; ou=ovule; ov=ovary; pe=petal; pr=parenchyma; se=sepal; st=staminal tube;
tt=transmitting tissue; vb=vascular bundle)
The ovules were anatropous and bitegmic and had
several cellular layers between the nucellus
epidermis and the embryo sac (Figures 31 and 32).
The integuments were multi-layered and the
number of layers varied from three to eight in the
outer integument and three to seven in the inner
integument (Figure 31). Two synergids of
prolonged contour were observed in the embryo
sac, where its filiform apparatus and the two polar
nuclei were notable (Figure 33).
The ovules had an obturator of placental origin
(Figure 32). The pistillode of the male flowers had
a structure similar to the pistil of the female
flowers, but with abortive ovules (Figure 2).
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Morphology and Anatomy of the Flowers
391
Figure 26-29. 26-27 - Details of the epidermis in the young and mature stigma of T. pallida and T. catigua,
respectively; 28 – 29: Detail of the transmitting tissue of the solid and hollow style of T. elegans and T.catigua,
respectively. (tt=transmitting tissue)
DISCUSSION
Pennington (1981) and Klein (1984) recorded for
Trichilia generally unisexual flowers in dioecious
plants and less frequently bisexual and then
polygamous plants. The flowers of T. catigua, T.
elegans and T. pallida were unisexual, but strictly
dioecious specimens were not found in the studied
area, all of them being monoecious to a greater or
lesser degree. The nectary presence in Trichilia
species in the form of a thick ring around the
ovary seemed to be common (Pennington, 1981),
as verified in T. elegans and T. pallida; although
in the latter species it was fused to base of
staminal tube.
According to Agthe (1951), the sugar
concentration of the nectar is correlated with the
type of vascular tissue that reaches the
nectariferous tissue. In present study, the vascular
tissue of the nectaries of the Trichilia species
consisted only of phloem. It meant that the nectar
sugar contents of these species could be high. This
supposition was in accordance with Agthe (1951)
who found similar results in other species from
different families.
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Souza, Luiz A. de et al.
Figure 30-33. 30 - Detail of the ovary wall of T. elegans; 31: Longitudinal section of the ovule of T. elegans; 32:
Diagram of the longitudinal section of the ovule of the T. catigua; 33: Embryonic sac of T. elegans showing
synergid cells and polar nuclei. (de=dorsal epidermis; ve=ventral epidermis; ii=inner integument; mi=micropyle;
nu=nucellus; ob=obturator; oi=outer integument; pl=polar nuclei; sc=secretory cell; sn=synergid cells)
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Morphology and Anatomy of the Flowers
The gynoecium structure is very important in the
reproductive process of the studied species, from
the arrival of the pollen on the stigma until the
occurrence of the syngamy inside the ovule. The
receptive stigma surface of the species constituted
by unicellular or bicellular trichomes seems to fit
in Group III of Heslop-Harrinson and Shivanna
(1977). This group is characterized by papillae or
trichomes of reduced or medium extension. The
transmitting tissue of the style, where the pollen
tube was developed, was mainly epidermal and
limited to a small central cavity in T. catigua and
T. pallida; but it was solid at the base and apex of
the T. elegans style. This meant that the pollen
tube developed along the style on the transmitting
tissue surface in the first two species or penetrated
this tissue growing among the cells in T. elegans.
When in the apex region of the ovary, the septs
were interrupted, facilitating communication
between the locules. This could favor eusyncarpy,
in which the pollen tube could fertilize an ovule of
any carpel -- a process verified in the Rutales,
very physiologically important and with
evolutionary consequences (Carr and Carr, 1961).
Finally, the obturator was found to be in contact
with the transmitting tissue, which facilitated the
penetration of the pollen tube into the micropyle.
The anatropous and bitegmic ovules seemed to be
characteristic for Trichilia species, as recorded in
the literature (Beltrati, 1984; Beltrati and Brunini,
1988, Moscheta, 1995 and Boesewinkel, 1981).
According to Boesewinkel (1981), the ovule of T.
grandifolia Oliv. is crassinucellate. According to
Davis (1966), the use of the terms crassinucellate
and tenuinucellate have been generating a lot of
confusion in the botanical literature. Therefore,
she introduced pseudocrassinucellate. According
to her, crassinucellate should be limited to the
ovule, in which the archesporial cell divides
forming a primary parietal cell which does or does
not divide, while pseudocrassinucellate should be
used for ovules, where no the parietal cells are
formed; but the apical cells of the nucellar
epidermis undergo periclinal divisions giving rise
to a nucellar cap. Thus, an ontogenetic study of the
nucellar development of the species would be
393
necessary to be able to affirm if the ovules are
crassinucellate or pseudocrassinucellate.
The synergids of the studied species presented
their respective filiform apparatus well-defined at
the basal end of the cell. Kapil and Bhatnagar
(1981) present several filiform apparatus shapes
for several Angiosperm species and attribute
several functions to this structure, such as
absorption, nutrient synthesis and transport,
glandular nature and an important role in the
penetration and effusion of the pollen tube, with
the consequent diffusion of spermatic cells to the
egg and the central cell (polar nuclei).
In several taxa one of the synergids can survive
after the fertilization, becoming hypertrophic,
polyploid or multinucleate and to exercising a
haustorial function - absorbing, storing or
secreting nutrients necessary for the zygote and
the young pro embryo (Kapil and Bhatnagar,
1981). It seemed that in T. catigua the two
synergids degenerated. This is confirmed by the
analysis of Moscheta (1995).
RESUMO
As características morfológicas e anatômicas das
flores de Trichilia catigua A. Juss., T. elegans A.
Juss. e T. pallida Sw. (Meliaceae) são descritas no
presente trabalho, visando subsidiar futuros
estudos sobre reprodução e preservação destas
espécies nativas. A antese ocorreu de abril a
agosto em T. catigua, de janeiro a maio em T.
pallida e setembro a janeiro em T. elegans. As
sépalas e pétalas apresentavam epiderme papilosa
e pilosa e mesofilo parenquimático. As flores
masculinas possuiam pistilódio com óvulos
abortivos e tubo estaminal com número variável de
anteras tetrasporangiadas, cuja parede apresentava
epiderme papilosa, endotécio, duas camadas
médias e tapete secretor. As flores femininas
apresentavam anteródios e pistilo bi ou tricarpelar,
com ovário de estrutura simples, estilete oco ou
sólido com tecido transmissor central e estigma
com pêlos uni ou bicelulares. Os óvulos eram
anátropos e bitegumentados. Não foram
encontrados no ambiente de estudo exemplares das
espécies estritamente dióicos, sendo todos em
maior ou menor grau monóicos.
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Souza, Luiz A. de et al.
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Brazilian Archives of Biology and Technology
Received: October 31, 2000;
Revised: January 05, 2001;
Accepted: April 27, 2001.