Progesterone and oestrogen receptor immunoreactivity in the vagina of the immature ostrich,Struthio camelus

1. The presence of sperm storage tubules in the cranial, middle and caudal regions of the vagina in the immature ostrich was studied. In addition, progesterone and oestrogen receptor immunoreactivity in these regions was investigated. 2. Sperm storage tubules were observed only in birds with active ovaries. Progesterone receptor immunoreactivity was more intense in birds with active ovaries, whilst immunostaining for the oestrogen receptor was absent in birds with both active and inactive ovaries. 3. These results suggest that steroid hormones produced by the active ovary mediate the activation of the progesterone receptor.     

Morphological features of the luminal surface of the magnum in the sexually immature ostrich (Struthio camelus)

Summary Observations were made, using scanning electron microscopy, of the surface features of the magnum in the immature ostrich during periods of ovarian inactivity, activity and regression. In birds with inactive ovaries the luminal surface of the magnum was lined with non-ciliated cells, which were densely covered by microvilli. In contrast, the magnum in birds with active ovaries was composed of ciliated and non-ciliated cells. The distribution of ciliated cells was not uniform, with clumps of cilia occurring next to non-ciliated areas. Samples collected from birds with regressing ovaries, during periods of decreasing daylength, revealed that the magnum was undergoing involution. The deciliation of ciliated cells and the presence of short microvilli on non-ciliated cells characterized magnal regression. These results suggest that ovarian activity and changes in daylength have a profound effect on the surface features of the magnum in the immature ostrich.     

Immunohistochemical localization of the progesterone and oestrogen receptors in the shell gland of sexually immature ostriches (Struthio camelus) with active or inactive ovaries

The immunohistochemical localization of progesterone and oestrogen receptors was studied in the shell gland of the immature ostrich (Struthio camelus) during periods of ovarian activity and inactivity. In birds with active ovaries moderate to strong immunostaining for the progesterone receptor was observed in the surface epithelium and tubular glands. In contrast faint progesterone receptor immunostaining was observed in the surface epithelium of the shell gland in ostriches with inactive ovaries. In addition, bud-like invaginations of the surface epithelium, which signaled tubular gland development, were negative for the progesterone receptor. Oestrogen receptor immunostaining, which was seen only in birds with active ovaries, was weak and restricted to nuclei of the surface epithelium. These results suggest that steroid hormones secreted by the active ovary regulate the differentiation of the shell gland. Furthermore, the influence of these hormones on the shell gland appears to be mediated predominantly through the activation of the progesterone receptor.     

A morphological and immunohistochemical study of healthy and atretic follicles in the ovary of the sexually immature ostrich (Struthio camelus)

The morphology of healthy and atretic follicles in the ovary of the sexually immature ostrich was described in the present study. In addition, the distribution of the intermediate filaments desmin, vimentin and smooth muscle actin, in these ovarian follicles, was demonstrated. Healthy and atretic primordial, pre-vitellogenic and vitellogenic follicles were present in the ovaries of the sexually immature ostrich. Atresia occurred during all stages of follicular development. Atretic primordial and pre-vitellogenic follicles were characterized by the presence of a shrunken oocyte surrounded by a multilayered granulosa cell layer. Two forms of atresia (types 1 and 2) were identified in vitellogenic follicles. In the advanced stages of type 1 atresia the follicle was dominated by a hyalinized mass. In contrast, in type 2 atresia the granulosa and theca interna cells differentiated into interstitial gland cells. Positive immunostaining for desmin was observed in the granulosa cells of only healthy primordial and pre-vitellogenic follicles. Atretic primordial and pre-vitellogenic follicles were immunonegative for desmin. Vimentin immunoreactivity was demonstrated in the granulosa cells of all follicles except the vitellogenic atretic follicles. The results of the present study indicate that ovarian follicles in the sexually immature ostrich undergo a cycle of growth and regression, which is similar to that reported in other avian species. Furthermore, based on the results of the immunohistochemical study, it would appear that the distribution and immunostaining of intermediate filaments changes during follicular development and atresia.     

Ultrastructural features of atretic follicles in the sexually immature ostrich (Struthio camelus)

The aim of this study was to describe the ultrastructural features of atresia in follicles of the immature ostrich (12–14 months old); a ratite that displays seasonal, precocious ovarian activity. The early stage of atresia in primordial, pre-vitellogenic and vitellogenic follicles was characterized by the accumulation of lipid droplets in the granulosa cells. Granulosa cells with condensed cytoplasm and nuclei were a prominent feature during the intermediate phase of atresia. The degenerating follicles were then infiltrated by stroma during the terminal stages of atresia. The results of this study provide further information on the morphology of atretic ovarian follicles in the immature ostrich.     

Macro-microscopic study on the toepad of ostrich (Struthio camelus)

The ostrich foot has four toepads, two on the 3rd digit, one on the 4th digit and one at metatarso-phalangeal joint. Previous studies have not detailed the histo-morphological structure of these toepads. In this study, we have described the macroscopic and microscopic structures of the toepad of ostrich (Struthio camelus). Numerous papillae with different direction, length and thickness have been observed grossly on the ventral surface of each toepad. Histological examinations have revealed that the epidermis of the ostrich toepad, similar to other digitigrades, consists of an outer stratum corneum and an inner stratum germinativum (which is subdivided into basal, intermediate and transitional layers). The stratum corneum has several layers of flattened horny cells. The nuclei of basal cells have several mitotic figures. The cytoplasm of the stratum germinativum cells has multiple lipid droplets and multigranular bodies (in transitional cells only). Scanning electron microscopic examination revealed presence of collagen fibers in mid and deep dermis of each toepad. These fibers run parallel and connect to each other by very thin fibrils which are branched, crossed with each other in an oblique direction. Such arrangement of these collagen fibers, thin fibrils and presence of digital cushion are likely to be responsible for the protection of the underlying soft tissues and absorption of concussion.     

A morphological study of simple testicular cysts in the ostrich (Struthio camelus)

Simple testicular cysts are rare in mammals and of unknown aetiology, but dietary conditions have been implicated in this phenomenon in poultry. This study characterises the structural features of simple intratesticular cysts in the ostrich. Seven testes from a total of 15 sexually mature ostriches slaughtered during the breeding season were used. The testes contained multifocal, fluid-filled, unilocular cysts which were lined by a simple squamous or low cuboidal epithelium and surrounded by a thick layer of fibrous connective tissue. The cysts developed within seminiferous tubules and displayed morphological features similar to those reported in man and domestic poultry. The testis parenchyma revealed several foci composed of intermingled normal seminiferous tubules and variably sized intratesticular cysts. The atrophic tubules lay within a mass of hyperplastic, fibroblastic intertubular connective tissue in the proximity of large cysts and their formation appeared to result from hydrostatic pressure exerted by cysts. Morphological evidence supports a continuous process of cyst formation in the affected testis and a concomitant progressive loss of atrophic seminiferous tubules. A pathogenetic scenario of cyst formation and the effect of simple cysts on testicular histology has been proposed. Although the course of this phenomenon remains unknown, its impact on the fertility of this economically important bird deserves closer scrutiny.     

Structural and immunohistochemical features of the epididymal duct unit of the ostrich (Struthio camelus)

The epididymal duct unit, comprising the ductus conjugens, ductus epididymidis and ductus deferens, was studied histologically, ultrastructurally and immunohistochemically in five sexually mature and active birds. The main morphological features of the pre-dominant non-ciliated (type III) cell of the epithelial lining of this duct unit include, but are not limited to, a moderately abundant smooth or sparsely granulated endoplasmic reticulum, electron-dense secretory granules and numerous mitochondria in the supranuclear zone of the cytoplasm. A single, large heterogeneous lipid droplet, of unknown function, was characteristically situated immediately proximal to the nucleus. The epithelium is obviously secretory and specifically, of the merocrine, and not apocrine, type of secretion. The epithelium of the epididymal duct unit was only focally and weakly to moderately immunopositive to both actin MF and desmin IF, while the duct unit was immunonegative to cytokeratin and vimentin intermediate filaments. The peritubular muscular layer was moderately to strongly positive to both actin and desmin, and negative to cytokeratins and vimentin.     

An ultrastructural characterization of the ooplasm in ovarian follicles of the immature ostrich (Struthio camelus)

Primordial, pre-vitellogenic and vitellogenic follicles were present in the ovary of the immature ostrich. The oocytes of these follicles were composed of a nucleus surrounded by ooplasm. Central, intermediate and cortical regions formed the ooplasm. The organelles present in these ooplasmic regions varied depending on the stage of follicular development. In primordial and small pre-vitellogenic (100-150 microm in diameter) follicles the central region of the ooplasm was dominated by an accumulation of organelles, which formed Balbiani's vitelline body. In contrast, the central region in vitellogenic follicles was filled with numerous large yolk spheres, many of which contained lining bodies. Numerous lipid droplets interspersed with mitochondria and small yolk spheres formed the intermediate ooplasmic region in primordial and small pre-vitellogenic follicles. In large pre-vitellogenic (150-400 microm in diameter) and vitellogenic follicles the intermediate region contained a greater density of mitochondria and small yolk spheres. Small yolk spheres were observed in the cortical region of pre-vitellogenic follicles. An interesting feature of the cortical region in vitellogenic follicles was the frequent occurrence of Golgi complexes. The results of the study indicate that although the ovarian follicles in the immature ostrich are not ovulated, the components and composition of the ooplasm are similar to those observed in the mature follicles of other avian species.     

Ultrastructural features of the follicular wall in developing follicles of the sexually immature ostrich (Struthio camelus)

The ultrastructure of the follicular wall in primordial, previtellogenic and vitellogenic follicles of the sexually immature ostrich is described in the present study. The follicular wall consists of a zona radiata, granulosa cell layer, basal lamina and thecal layer. Cytoplasmic processes from the plasma membranes of the granulosa cell layer and the ovocyte form the zona radiata in previtellogenic and vitellogenic follicles. The granulosa cell layer transforms from simple cuboidal epithelium in primordial follicles to simple columnar or pseudostratified columnar epithelium in previtellogenic and vitellogenic follicles. Transosomes were observed along the apical and lateral plasma membranes of granulosa cells. The thecal layer in previtellogenic and vitellogenic follicles consists of interna and externa components. The fibroblasts in the theca externa contain microfilaments, which are thought to be actin filaments. The study revealed ultrastructural features, which are associated with the transportation of yolk precursors and nutrients into the ovoplasm. In addition, the study indicates that, although the cells in the theca externa contain microfilaments, they do not possess the ultrastructural characteristics of smooth muscle cells.     

Growth in the ostrich (Struthio camelus var. domesticus)

The use of the ostrich as a livestock animal necessitates an understanding of its ability to utilize specific nutrient requirements if efficient growth is to be realized. Male ostriches may have to be raised separately as they grow faster; they require higher protein diets and are more efficient feed converters for longer periods than female ostriches. Male ostriches exhibit slightly faster maturation rates than female ostriches. The corresponding age at which maximum growth is attained is 181 days for male ostriches and 199 days for female ostriches. The growth curve of ostriches is sigmoid and asymptotic. Body mass reached in 1 year is approximately 104 kg. The metabolic age of ostriches at hatching is approximately 13.7 days. Feed conversion has an impact on growth. The ostrich chick (aged <2 months) has a high feed conversion to body mass ratio of 2:1, which requires very accurate and careful management of chick nutrition.     

A light microscopic and immunocytochemical study of the gastrointestinal tract of the ostrich (Struthio camelus L.)

Although the histological features and endocrine cells of the gastro-intestinal tract of the chicken have been well studied, little is known about these features of the gut of the ostrich. The present study was undertaken to elucidate the histology and peptide-storing endocrine cells of the ostrich. As a rule the histological features of the gastro-intestinal tract of the ostrich corresponded to that of the fowl. However, certain differences were observed. The superficial proventricular glands were simple, branched tubular glands, while the deep proventricular glands were restricted to a slipper-shaped area and extended into the muscularis mucosae. The gizzard had a variably developed muscularis mucosae, a feature that seems to be unique to the ostrich. The villi of the small intestine were long and branched profusely, forming a labyrinthine surface. No Paneth cells were observed. The mucosa of the ceca and the first part of the rectum was thrown in large circular folds, forming a compressed spiral. Numerous melanocytes were seen in the submucosa and the connective tissue around the bloodvessels of the muscle layers at the tips of the ceca. A well developed subserosa was present throughout the gastro-intestinal tract. Endocrine cells immunoreactive to somatostatin, glucagon, gastrin, bombesin, neurotensin, substance P and pancreatic polypeptide were detected in the gastro-intestinal tract of the ostrich. The topographical distribution of those endocrine cells immunoreactive to glucagon, bombesin, neurotensin and substance P differed from that of the chicken. The results of this investigation inferred that at least one of the gut peptides of the ostrich (secretin) to be structurally different from its counterparts in mammal and chicken. Molecular heterogeneity of somatostatin was observed in endocrine cells situated in the deep ventricular glands of the ostrich.     

Development of the digestive tract in the ostrich (Struthio camelus).

A study was conducted to investigate changes in the development of the gastrointestinal tract (GIT) in relation to body growth of growing ostriches. There was an 11-fold increase (P < 0.001) in body weight between 3 and 72 days of age. The relative (to body weight) weight of the proventriculus/gizzard, caeca and colon also increased (P < 0.001) with age. The relative weight of the small intestine peaked at 41 days of age and then tended to decline (P < 0.05) subsequently. The relative weight of the pancreas peaked at 27 days of age and remained fairly stable thereafter. The activities of chymotrypsin and lipase declined (P < 0.001) with age between 3 and 72 days. At 3 days of age, the protein content of the duodenal mucosal homogenate was higher (P < 0.001) than that of the jejunum or ileum, but at all subsequent periods the jejunal protein content was the highest. The protein content of the intestinal brush-border membrane was higher (P < 0.001) at the jejunum than at the duodenum or ileum. The specific activity of maltase declined (P < 0.001) with age in all three regions, most especially between 3 and 27 days of age. The activity of alkaline phosphatase (AP) at 41 and 55 days of age was higher (P < 0.001) in the duodenum than in the jejunum or ileum. The activity of AP fluctuated with age in the duodenum but there was a more defined decline (P < 0.001) with age in the jejunum and ileum. The relative protein content of the liver increased (P < 0.001) with age, with two peaks at 27 and 55 days of age. Arginase activity was not detected in the liver of 3-day old chicks and was not significantly affected by age between 27 and 72 days of age. The pattern of development observed is similar to that in growing poultry. There is, however, a need for evaluation at closer intervals in early life as well as an in-depth assessment of the morphometry of the intestinal mucosa.     

A case of conjoined cephalopagus twinning in an ostrich (Struthio camelus)

Conjoined twinnings have been reported in most domestic animal species and in some avian species. Cases of conjoined twins have not been reported in the ostrich so far. A hybrid Blue neck x African black male ostrich conjoined twinning was born at the end of artificial egg incubation and died spontaneously 24 h after the hatching. It was frozen and sent to the Unit of Veterinary Pathology of the University of Messina for gross examination. The most important gross findings involved the external body and most of the internal organs. On the basis of the duplication, the conjoined twins were classified as a cephalopagus. Radiological features included: development of one head containing a single brain, two spinal cords, deviated vertebral columnae with fusion of the two first cervical vertebrae. In one twin, the synsacrum was absent as well as portions of the vertebral column. Grossly, both twins showed two upper and lower limbs each. The gastro-enteric apparatuses of the twins were not completely developed and fused at different levels. One liver and one heart localized in the centre of the conjoined twins were observed. The authors conclude that the possible causes of the malformation could be related to a genetic factor.     

Characterisation of tetra-nucleotide microsatellite loci in the ostrich (Struthio camelus)

1. Eleven polymorphic tetra-nucleotide microsatellite loci were identified in the ostrich (Struthio camelus) using a selective enrichment protocol.

2. The average number of alleles observed was 6·6 with an average heterozygosity of 0·4.

3. The population was found to be in Hardy-Weinberg equilibrium and two of the loci had a greater than 95% probability of having null alleles.

4. These microsatellite loci will add to the existing pool of markers available for the ostrich and help to facilitate analysis of population structure and pedigree determination.     

The topography of the thoraco-abdominal viscera in the ostrich (Struthio camelus)

The topography of the thoraco-abdominal viscera in the ostrich was studied in 20 birds varying in age from 2 weeks to 12 months. The lungs occupied the dorsal third of the thorax, and the heart lay in the cranioventral thorax perpendicular to the long axis of the body. There was no pleural cavity. The liver was situated in the caudoventral part of the thorax, and the proventriculus occupied the left cranial part of the abdomen between the 7th vertebral rib and the acetabulum. The gizzard lay in the cranioventral part of the abdomen, resting on the sternum and abdominal floor. The duodenum formed a loop from right to left, with the pancreas lying between the 2 limbs of the loop. The coiled jejunum and ileum occupied the ventral part of the abdomen between the gizzard and pelvis. The two caeca lay on either side of the terminal ileum with their apices in the pelvis. The rectum was the longest part of the intestine and could be divided into a thick proximal segment situated in the right dorsal part of the abdomen, and a thin distal part that occupied the left caudodorsal part of the abdomen. The trilobed kidneys lay along the ventral surface of the synsacrum, with the adrenal glands at their cranioventral poles. The testes lay ventrally to the cranial divisions of the kidneys, whereas the left ovary was situated ventrally to the cranial division of the left kidney. The spleen lay wedged in between the right kidney, caudal vena cava and proventriculus. The thyroid glands were situated at the cranial borders of the subclavian arteries, and the thymus lay at the base of the neck.     

Radiographic anatomy of the thoraco-abdominal cavity of the ostrich (Struthio camelus)

This study provides a reference for the radiographic anatomy of the thoraco-abdominal cavity of female ostriches as a representative of ratites. One ostrich cadaver, 2 adult and 2 growing ostriches were used. Right lateral radiographs produced by a 6-frame technique and 2 dorsoventral radiographs produced by an adapted 3-frame technique were selected and schematic illustrations of these were labelled to illustrate normal radiographic anatomy. Differences from other avian species and unique features of the ostrich are briefly discussed.     

Ultrastructural features of the uterus in the sexually immature ostrich (Struthio camelus) during periods of ovarian inactivity and activity

The ultrastructure of the surface epithelium and tubular glands of the uterus in the immature ostrich is described. In ostriches with inactive ovaries the uterus is lined by a non-ciliated simple columnar epithelium, with basally located heterochromatic nuclei. Scanning electron microscopy revealed that these non-ciliated cells have a dense microvillous cover. A simple columnar to pseudostratified columnar epithelium, comprised of non-ciliated and ciliated cells, lines the uterus in birds with active ovaries. The ciliated cells possess a wide luminal region, which contains a nucleus and various organelles. An accumulation of secretory granules was observed in the apical regions of the non-ciliated cells, as well as in a few ciliated cells. In addition to non-ciliated and ciliated cells, a cell type with rarefied cytoplasm was also identified. These cells appear to correspond to calcium secreting cells identified in other avian species. The results of this study indicate that, although uterine differentiation is present in immature ostriches with active ovaries, the production of secretory product appears to occur mainly in non-ciliated epithelial cells.     

Feeding and pecking behaviour in ostrich (Struthio camelus) chicks in captivity

1. Three sets of experiments were performed on two batches of ostrich chicks to investigate the factors affecting the pecking and feeding behaviour of grouped individuals.

2. Chicks showed no significant alteration of their feeding behaviour in response to raising pen walls in the rearing facility from 30 to 60 cm.

3. Further analysis on a different set of birds revealed consistent short term individual differences in the frequency of feeding and non‐feeding pecks.

4. There were significant pen effects on behaviour suggesting the possible development of a pen ‘culture’ of pecking behaviour.

5. Pecking behaviours in 26 to 33 d‐old chicks, with the exception of drinking, were generally negatively correlated, so any non‐food pecking by a chick was generally associated with fewer pecks targeted at food.

6. By the age of two months chicks were pecking at food on the floor to a far greater extent than at any food presented in food trays.