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Revista chilena de anatomía

versión impresa ISSN 0716-9868

Rev. chil. anat. v.15 n.1 Temuco  1997 



* Catroxo, M. H. B.
** Lima, M. A. I.
* Cappellaro, C. E. M. P. D. M.

* Instituto Biológico de São Paulo, São Paulo -Brazil.
** Department of Morphology - Histology - Universidade Federal de São Paulo/ Escola Paulista de Medicina, São Paulo, Brazil.

SUMMARY: A morphological (histological) study of the stomach (proventriculus and gizzard) of the red-capped cardinal (Paroaria gularis gularis) was carried out, under light microscopy. Anatomically, the stomach of the red-capped cardinal is constituted by two externally distinguishable chambers: a cranial region, glandular, the proventriculus (proventriculus, pars glandularis), which is cranially connected to the esophagus and caudally, to the gizzard or ventriculus (ventriculus, pars muscularis), also known as muscular portion. Both, the mucous tunic of the proventriculus and of the gizzard present folds lined by simple prismatic epithelium. A thick cuticle is laid over the mucous tunic of the gizzard. In the lamina propria of both regions, there are simple tubular glands. The submucosa of the proventriculus is occupied by deep proventriculus glands. Due to the absence of a muscularis mucosae, the submucosa of the gizzard cannot be distinguished from the lamina propria. The muscle tunic of proventriculus, consists of a inner longitudinal layer, of an intermediary circular layer and of an external longitudinal discontinuous layer of smooth muscle. In the gizzard, it consists of an internal longitudinal layer and of an external circular layer. In both chambers, the serosa is constituted by a connective tissue lined by mesothelium, containing blood vessels, nerve elements of the serous plexus and adipose tissue.

KEY WORDS: 1. Birds; 2. Proventriculus; 3. Gizzard; 4. Digestive system; 5. Histology.


The stomach (gaster) structure of the birds presents variations which depends on the alimentary habits of the bird (FARNER, 1960; STURKIE, 1965 and McLELLAND, 1979).

The stomach of the red-capped cardinal, as well as in other birds, consists of two externally distinguishable chambers: the proventriculus (proventriculus) or glandular part (pars glandularis) which secretes the gastric juices, and the gizzard (ventriculus) or muscular part (pars muscularis) which has a mechanical function (FARNER; McLELLAND and BRADLEY & GRAHAME, 1951).

The proventriculus is cranially continuous with the esophagus and presents an eliptical shape. The gizzard or ventriculus (ventriculus) extends caudally to the proventriculus and is round and flat-shapped with a very thick wall.

With the purpose to correlate the morphology of the stomach with the alimentary habits of birds in general, we carried out a histological study of the stomach (proventriculus and gizzard) of the red-capped cardinal (Paroaria g.gularis), which basic alimentary habits are composed by seeds, pulp of fruit and insects.


Four birds (males, adults) were captured in the city of Peruíbe, SP, Brazil and kept in individual cages, for a short period. They were fed with grains and fruits similar to those existent in the area of capture and "ad libitum" water. After anesthesia with ether inhalation, the digestive tube was exposed and fragments of the stomach (proventriculus and gizzard) were immediately fixed in Bouin’s liquid, for 24 hours. After dehydratation with ethylic alcohol in increasing concentrations (70 to 100%) the fragments were embedded in paraffin, sectioned at 5 µm and stained by haematoxylin-eosin and by the CASTRO & CAMARGO (1951) polichromic stain. The nominal description of the regions were according to Nomina Anatomica Avium (Mc LELLAND, 1979).


The analysis of the histological sections, under light microscopy, revealed that the wall of the stomach (proventriculus and gizzard) is constituted by the following layers: mucous membrane (tunica mucosa gastris), submucosa (tela submucosa gastris), muscular (tunica muscularis gastris) and serosa (tunica serosa).

The mucous membrane (tunica mucosa gastris) of proventriculus (proventriculus) presents folds (plicae proventriculares) in its luminae surface, lined by a simple prismatic epithelium (Fig.1). The lamina propria is constituted by connective tissue (Fig.2) with blood vessels and lymphocitary infiltration. In its interior, simple tubular glands are found (gll. proventricularis superficiales), lined by the same type of epithelium of the surface (Fig.2). Isolated muscular smooth fibers, located together the lamina propria, constitute a incipient muscularis mucosae (lamina muscularis mucosae). The submucosa is constituted by connective tissue, containing blood vessels. The deep proventricular glands are located in it (gll. proventriculares profundae). Those conic or pear-shapped glands occupy great part of the thickness of the wall of the proventriculus. Each gland presents a connective tissue capsule and is constituted by numerous secretory tubules lined by cuboid cells, with deep stained eosinophilic cytoplasm. These cells are juxtaposed in its basal portions, providing to the epithelium a serrated appearance. Covering each secretory tubule, we observed a delicate connective tissue fillet (Fig.2). Each secretory tubule continues by one duct which opens in the main collecting duct, which leads to the luminal surface of the organ.

In those composed tubular glands, the ducts are lined by prismatic cells with clear cytoplasm (Fig.3). The muscle tunic (tunica muscularis gastris), presents an inner layer of longitudinaly disposed smooth muscular fibers (stratum longitudinale) and another intermediary, of circulary disposed fibers (stratum circulare) (Fig.1). Externally, we observed a discontinuous layer, constituted by bands of longitudinally arranged smooth muscular tissue (stratum longitudinale). Among the most external bundles of muscular fibers, we observed nerves and nervous ganglions of the mioenteric or of the Auerbach plexus. The serosa of the proventriculus (tunica serosa) is constituted by connective tissue lined by mesothelium, and is rich in blood vessels and nervous elements (nerves and ganglions) of the serose plexus (Fig.1).

The mucous membrane (tunica mucosa gastris) of the gizzard presents low folds (plicae ventriculares), which are lined by simple prismatic epithelium (Fig.4). Over the mucous membrane, a thick cuticle is disposed (cuticula gastrica) (Fig.4). The lamina propria is constituted by a dense connective tissue and it is occupied by numerous deep simple tubular glands (gll. ventriculares) which expand in the base of the fold, paralelly located between them (Fig.4). Those glands are lined by a simple epithelium, which is lower in the base of the glands and higher in their upper portion (Fig.4). The cells of the glands present nucleus with very improved nucleolus. In the interior of the glands, we observed eosiniphilic secretion fillets, continuous with the cuticle (Fig.4). Some glandular tubes are narrower, other wider. We have not identified the muscularis mucosae (lamina muscularis mucosae). Due to lackness of the muscularis mucosae, we could not identify the submucosa layer (tela submucosae gastris). The muscle tunic (tunica muscularis gastris) (Fig.4) is constituted by a longitudinal layer (stratum longitudinale) and by a developed outer circular layer (stratum circulare) of the smooth muscle. The muscular bundles are interpersed with bands of connective tissue (Fig.4), rich in blood vessels and nervous ganglions of the mioenteric or of the Auerbach plexus. The serosa (tunica serosa), which is constituted by connective tissue lined by mesothelium, is rich in blood vessels, nervous elements of serous plexus and adipose cells.


Our histological results disclosed that the structure of the proventriculus and of the gizzard are similar to those of other birds, according to what is described by several authors (McLELLAND; CALHOUN, 1954; HODGES, 1974 and FIERI, 1984). So, we could observe that the mucous membrane of the proventriculus and of the gizzard present several folds,

Fig 1 - Photomicrography of the cross section of the proventriculus: folds of mucous membrane (P);
deep proventricular glands (GP); capsule of gland (arrow head); muscle tunic (m); serosa with blood
vessels (S); lumen of the organ (L). Hematoxylin-eosin stain. X50.



Fig. 2 - Photomicrography of longitudinally section of the proventriculus showing: simple tubular glands (Gs) in the lamina propria (Lp), constituted by a simple prismatic epithelium; part of deep proventricular gland (GP), constituted by secretory tubules (T), lined by cuboidal cells
(c). Observe (arrow head) thin bundles of connective tissue around the deep glands. Castro & Camargo polychromic stain. x 300.



as in other birds (McLELLAND; BRADLEY & GRAHAME; CALHOUN and FIERI) the folds of the gizzard are lower. In the proventriculus, the shape and arrangement of those folds vary as branched (LIMA, 1979 and LIMA & SASSO, 1985) anastomotic (KLEM et al., 1982; KLEM et al., 1984 and ROCHA, 1991), or constituted by foliaceous vilosities (FIERI). In the gizzard, among several kinds of birds, the folds of the mucosa are described as longitudinal (JAIN, 1976 in P. krameri; MENIN et al., 1990 in Coragyps atratus foetens), and high (LIMA, 1979 in Columba livia) or parallel among (ROCHA, in Speotyto cunicularia).

Fig. 3 - Photomicrography of the cross section of the proventriculus presenting: capsule of connective
tissue (arrow head) around the glands (GP), constituted by numerous secretory tubules (T). Each
secretory tubule continues by one duct (d), which opens in the main collecting duct (D), which leads to
the luminal surface of the organ (L). Castro & Camargo polychromic stain. x 160.


Fig. 4 - Photomicrography of longitudinally section of the gizzard showing: folds of mucous membrane lined by simple prismatic epithelium (P); simple tubular glands (Gs) in the lamina propria constituted by connective tissue (Lp). Observe: secretion fillets (S), continuous with the cuticle (C), part of muscle tunic red stained (m), interpersed with bundles of connective tissue (Tc). Castro & Camargo polychromic stain. x 210.






KLEM et al. (1983, 1984) described the mucous membrane of the gizzard of Passer domesticus and of Turdus migratorius as constituted by dual viliform folds, whereas VITTORIA & RICHETTI, 1974 described it in the gizzard of carnivorous and granivorous birds, as composed by eliptical or circular crypts which unfolds as to the surface. AKESTER, 1986 stated that in Gallus gallus the eliptical crypts may reach the shape of large and narrow fissures. The lining epithelium of the folds of the mucous membrane of the proventriculus is of prismatic type, which coincides with the observations of most of the authors (STURKIE; CALHOUN; HODGES; FIERI; LIMA, 1979 and 1985; KLEM et al., 1982 and 1984; ROCHA; SELANDER, 1963; SINGH, 1973; VIAL et al., 1977 and BEE DE SPERONI & CHIKILIAN, 1983. The lining epithelium of the folds of the gizzard is of simple prismatic type. Similar to our observations GLEREAN & KATCHBURIAN describe a high prismatic type epithelium, lining the mucous membrane of the gizzard of the Gallus gallus.

In the lamina propria of both chambers, we have found numerous simple tubular glands, which open in the base of the folds. Those results coincide to those of FARNER in birds in general, with to the LIMA , in Columba livia and with to the ROCHA in Speotyto cunicularia. The localization of those glands, in the proventriculus, varies among the several kinds of birds. ANDREW (1959) describes their presence, in birds in general, between the ducts of the composed glands; VIAL et al. describe them in the luminal surface of the mucosa of Columba livia and FIERI has found them in the base of the vilosities of the proventriculus of Nothura maculosa maculosa. Similarly to our conclusions, some authors have found the proventriculus simple tubular glands lined by simple columnar epithelium (FIERI; ROCHA and ESPINOLA & GALLIUSSI, 1990). FARNER describe them, in birds in general, as lined by cuboid-type epithelium. In the lining of the simple tubular glands of the gizzard, we confirmed the predominance of prismatic cells, lower in the base of the gland and higher in their upper region. Those results are similar to those of some authors (GLEREAN & CASTRO, 1965 in Thryothorus longirostrus, FIERI in Nothura maculosa maculosa and ESPINOLA & GALLIUSSI in Fulica armillata). On the other hand, other authors have described such epithelium as constituted by cubic to prismatic cells, in Gallus gallus (GLEREAN & KATCHBURIAN, 1964), as varying from cuboid to columnar epithelial cells in birds in general (STURKIE) and in Gallus gallus (EGLITIS & KNOUFF, 1962); KLEM et al. 1983 and 1984, in Passer domesticus and Turdus migratorius; as cubic cells, in birds in general (BRADLEY & GRAHAME; CALHOUN and LOBO et al., 1960) in Coragyps atratus foetens (JAIN) in P. krameri (LIMA, 1979), in Columba livia and in Milvago chimachima chimachima (GLEREAN & CASTRO).

As far as the distribution of the simple tubular glands of the gizzard is concerned, we have observed them as paralelly arranged among them, presenting some glandular narrow and other wider tubes. In the inner part of those glands, we have observed fillets of eosinophilic secretion, along with the cuticle, which was also observed by LIMA who described in the lumen of the glands of the gizzard in Columbia livia, an haematoxylin-eosine intensive red-stained secretion, going towards the surface as columns, so as to merge to the thick covering plate. Lining the mucous membrane of the gizzard we have identified a thick layer, constituting the cuticle. Those conclusions are similar to those statements in Literature which also refers to the mucosa surface cover as cuticle (Mc LELLAND; FIERI; KLEM et al., 1983 and 1984; ROCHA and ANDREW), although some refered to have such layer as "Koilin Membrane"(AKESTER; TONER, 1964 and LANDOLT, 1985), formed by the secretion of the gland, by the crypt and by the surface lining epithelial cells (AKESTER; EGLITIS & KNOUFF; TONER and LANDOLT). As far as the aspect of the lamina propria of the proventriculus and of the gizzard is concerned, our conclusions are similar to those already discussed as constituted by connective tissue (CALHOUN, FIERI; KLEM et al.,1982, 1983 and 1984 and EGLITIS & KNOUFF), rich in blood vessels (CALHOUN, FIERI, KLEM et al., 1982 and 1984 and ESPINOLA & GALLIUSSI), lymphatic (ESPINOLA & GALLIUSSI) and limphocitary elements in some places, in the proventriculus (HODGES and FIERI). We have found isolated smooth muscular fibers closed to the lamina propria of the proventriculus, which indicates a muscularis mucosae, also mentioned by ROCHA in the muscularis mucosae layer of the proventriculus of Speotyto cunicularia. Such layer, also described by ANDREW in the proventriculus of birds in general is constituted by an inner circular and an external longitudinal layer of smooth muscular fibers, which FEDER (1969) and BEE DE SPERONI & CHIKILIAN have observed as extremelly thin in the proventriculus of Melosittacus undulatus, Zenaida auriculata chrysaychenia and Myiopsitta monacha monacha, respectivelly.

In the gizzard, we have not identified the musculares mucosae which is accordingly to the statements of most of authors (Mc LELLAND; CALHOUN; FIERI; ANDREW and EGLITIS & KNOUFF) . On the other hand, ROCHA has detected such layer, much developed in Speotyto cunicularia (carnivorous bird), constituted by longitudinally disposed smooth muscular fibers, located under the lamina propria. On the other hand, JAIN has described, as in the gizzard of P. krameri (frugivorous bird) a poorly developed muscularis mucosae which is absent in L. schach (carnivorous bird) and in A. tristis (omnivorous bird).

Our observations, regarding the submucosa of the proventriculus, correspond to the most of the researches, who have described it as constituted by connective tissue (FARNER; Mc LELLAND; BRADLEY & GRAHAME; HODGES; FIERI; KLEM et al., 1982 and 1984 and ROCHA). Others have described it as extremelly thin Mc LELLAND , in birds in general, or thin in some places and not present in others (CALHOUN, in chicken, HODGES , in birds in general and FIERI, in Nothura maculosa maculosa) or poorly developed (FARNER , in birds in general). We have found a numerous blood vessels in the submucosa of proventriculus, as well as JAIN in L. schach (carnivorous bird) and SINGH in M. haemacephala (frugivorous bird). Regarding the deep proventricular glands, we have found them in this tunic, under the isolated smooth muscular fibers of the muscularis mucosae, which was also reported by other researchers (BRADLEY & GRAHAME; ROCHA; SELANDER; SINGH; LOBO et al.; TONER and HELANDER, 1981). In the red-capped cardinal, those glands are tubular composed, as also mentioned by other authors (KLEM et al., 1982 and 1984; VIAL et al.; BEE DE SPERONI & CHIKILIAN; LOBO et al.). Others have found them as branched tubular glands (FIERI, LIMA & SASSO, 1985 and ROCHA). The shape of the glands, according to our studies, is similar to those described by LIMA and VIAL et al. in Columba livia and FIERI in Nothura m. maculosa, as being conical or pyriform structures. SINGH has described them in Cinnyris asiatica (nectiverous bird) as eliptical or long-shapped, whereas FARNER reported that, in birds in general, they may be irregular, cilindric, eliptical or circular. MARSHALL & FOLLEY (1956), have described in Collocalia brevirostris and in Collocallia francica (insectivorous bird) with each gland extending externally into a membranous tube which splits into separated portions like the petals of flowers. We have identified in those glands a connective tissue capsule which involve the numerous secretory tubes. Similarly to those conclusions, other authors also described thin layers of connective tissue, surrounding the deep of the proventriculus glands, containing many blood vessels and nerves ( BRADLEY & GRAHAME; CALHOUM; FIERI; LIMA; KLEM et al., 1982 and 1984; ROCHA; VIAL et al.; ESPINOLA & GALLIUSSI and FEDER). The system of collecting ducts which we have found coincides,in general aspects, to what has been already described by several authors, in which the glandular unities are radially disposed around a central collecting chamber, from which a duct which communicates with the proventriculus emerges (FARNER; CALHOUN; HODGES; FIERI; VIAL et al.; LOBO et al.; ANDREW; ESPINOLA & GALLIUSSI; HILL; HELANDER and TURK, 1982). Similarly to our conclusions, many other authors describe a columnar or simple prismatic lining epithelium in the ducts (FARNER; BRADLEY & GRAHAME; CALHOUN; FIERI; LIMA; KLEM, 1982 and 1984; ROCHA; BEE DE SPERONI & CHIKILIAN; ESPINOLA GALLIUSSI and LANZIERI et al., 1983).

We found numerous secretory tubes lined by cuboid cells, similar to the observations of BRADLEY & GRAHAME; TONER; FEDER; SINGH; LIMA; FIERI; ROCHA. Other authors described the glandular epithelium as constituted by polygonal (KLEM et al., 1982 and 1984) or conic-shapped cells (LOBO et al.) or, further, varying from low cuboid to elongated columnar shape (HILL). Similarly to the results obtained by JAIN in L. schach (carnivorous bird) and by A. tristis (omnivorous bird) and by ROCHA in Speotyto cunicularia, we have not identified the submucosa in the gizzard. Some authors describe the submucosa as constituted by one thick connective tissue layer (Mc LELLAND; CALHOUN; FIERI and EGLITIS & KNOUFF), by a thin connective tissue strings (KLEM et al., 1983) or by a thin connective tissue layer (HILL). We have identified in the proventriculus, the muscle tunic constituted by three muscular fibers layers disposed in an internal longitudinal stratum, a medium circular stratum and a discontinued external circular stratum. Those observations are similar to those of BRADLEY & GRAHAME , in the muscle tunic of the proventriculus of birds in general. ESPINOLA & GALLIUSSI and BANKS (1991) have also observed those three layers of the muscle tunic in Fulica armillata and in birds in general, respectivelly, disposed in an inner longitudinal layer, a medium circular layer and an extenal discontinued longitudinal layer of smooth muscular fibers.

CALHOUN and ANDREW have described, in the proventriculus of birds in general, the muscle tunic as constituted by an external layer of smooth muscular fibers, longitudinally disposed and a circular arranged inner layer. Most of the authors reported that the inner circular layer is thicker than the external longitudinal one (HODGES; FIERI; ROCHA; BEE DE SPERONI & CHIKILIAN and FEDER). Regarding the presence of nerves and nervous ganglions of the miontheric plexus of Auerbach, we have found them among the bundles of most external smooth muscular fibers. FIERI and ROCHA refer to the presence of mioenteric nervous plexus (Auerbach) among the muscular layers of the proventriculus of Nothura maculosa maculosa and of Speotyto cunicularia, respectively. In the gizzard, the muscle tunic presents some variation regarding the number and the arrangement of its layers. In our research, we could find the muscle tunic as constituted by an inner layer of longitudinaly arranged smooth muscular fibers and an circular arranged external layer, much more developed. Regarding the arrangement of its layers, our results are opposite to those of some authors whose description are refered as constituted by an inner circular layer and one external longitudinal layer (JAIN, in P. krameri, frugivorous bird and in A. tristis, omnivorous bird; FIERI, in Nothura maculosa maculosa, (insectivorous and granivorous bird); ROCHA in Speotyto cunicularia (carnivorous bird); Mc LELLAND, and TURK, in birds in general. However, those same researchers also related that the circular layer is more developed if compared to the longitudinal one. Some observations in Literature refer to the presence, in several birds, of three layer in the muscle tunic, in different arrangements. ESPINOLA & GALLIUSSI have also described three layers in the muscle tunic of the gizzard of Fulica armillata (granivorous bird): one internal of very thin and discontinuous muscular fibers, one meddium sized, very thick, longitudinally disposed; and one external constituted by smooth muscular fibers which is thiner than the first one, circularly arranged. IMAIZUMI & HAMA (1969) , also related the existence of three muscular layers in the gizzard of the Uroloncha domestica, disposed in an inner obliquelayer, a medium sized layer and an external longitudinal layer. Considering what has been reported so far, we have observed that there are a great variation as far as the development of the muscle tunic among the several kinds of birds which were studied is concerned. AKESTER states that, in birds whose alimentary habits are restricted to grains and insects, the gizzard presents much developed muscles and, as the food is smooth as meat and fruits, the muscles are much less developed. In Pelecanus phillippensis (piscivorous bird), SINGANALLUR et al., (1976) associated the much developed and extremelly thick muscular of the gizzard to the function of triturating and crushing of the body of the fish. AKESTER stresses that, in one extreme situation, one observes that in some frugivorous birds, almost all the gizzard disappears. Confirming those facts, KOLB (1984) mentions that the relationship between the structure of the gizzard and the kind of food can be established by the atrophy of the muscles, as granivorous birds are fed with smooth food. The presence of much developed muscle tunic in the gizzard of the red-capped cardinal, which alimentary habit is based in grains, fruits and insects, is according to the relation existent between morphology and alimentary habits, established by the above mentioned authors. We have also observed the presence of connective tissue bundles mixed with the muscle bundles in the gizzard, as well as blood vessels and nervous ganglions of the mioenteric plexus (of Auerbach), among the muscular bundles, similar to those described by FIERI in Nothura maculosa maculosa and by ROCHA in Speotyto cunicularia. The description of the serosa of the proventriculus and of the gizzard of the red-capped cardinal corresponds to that of most of authors, as constituted by connective tissue, containing many blood vessels and neves, and nervous elements (nerves and ganglions) of the serous plexus (CALHOUN; FIERI and ROCHA) and adipose tissue (FIERI) lined by mesothelium (ROCHA).

RESUMEN: Un estudio morfológico (histológico) del estómago (proventrículo y molleja) del cardenal rojo (Paroaria gularis gularis) fue efectuado bajo microscopio de luz. Anatómicamente, el estómago del cardenal rojo está constituido por dos cámaras distintas: la región craneal, glandular o el proventrículo (proventriculus, pars glandularis), la cual se está conectada cranialmente con el esófago y caudalmente, está el ventrículo (ventriculus, pars muscularis) también conocida como porción muscular. Ambas, la túnica mucosa del proventrículo y del ventrículo presentan pliegues alineados de epitelio simple prismático. Una cutícula densa está colocada encima de la túnica mucosa del ventrículo. En la lámina propia de ambas regiones, se encuentran glándulas tubulares simples. La submucosa del proventrículo está ocupado por glándulas proventriculares profundas. Debido a la ausencia de una mucosa muscular, la submucosa del ventrículo no puede ser distinguido de la lámina propia. La túnica mucosa del proventrículo, consiste de un lecho interior longitudinal, de un lecho intermedio circular y de un lecho externo longitudinal discontinuo de músculo liso. En el ventrículo, consiste de un lecho longitudinal interno y de un lecho circular externo. En ambas cámaras, la serosa está constituida por tejido conectivo revestido por mesotelio, conteniendo vasos sanguíneos, elementos nerviosos del plexo seroso y tejido adiposo.

PALABRAS CLAVE: 1. Pájaros; 2. Proventrículo; 3. Molleja; 4. Sistema digestivo; 5. Histología


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Adress for corresponding:
Dra. Marcia Helena Braga Catroxo
Department of Electron Microscopy
Instituto Biológico
Av. Conselheiro Rodrigues Alves, 1252
CEP: 04014-002
São Paulo - S.P.

Recibido : 13-12-1996
Aceptado: 06-03-1997

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