The Harderian gland: Endocrine function and hormonal control.

The Harderian gland (HG) is an exocrine gland located within the eye socket in a variety of tetrapods. During the 1980s and 1990s the HG elicited great interest in the scientific community due to its morphological and functional complexity, and from a phylogenetic point of view. A comparative approach has contributed to a better understanding of its physiology. Whereas the chemical nature of its secretions (mucous, serous or lipids) varies between different groups of tetrapods, the lipids represent the more common component among different species. Indeed, besides being an accessory to lubricate the nictitating membrane, the lipids may have a pheromonal function. Porphyrins and melatonin secretion is a feature of the rodent HG. The porphyrins, being phototransducers, could modulate HG melatonin production. The melatonin synthesis suggests an involvement of the HG in the retinal-pineal axis. Finally, StAR protein and steroidogenic enzyme activities in the rat HG suggests that the gland contributes to steroid hormone synthesis. Over the past twenty years, much has become known on the hamster (Mesocricetus auratus) HG, unique among rodents in displaying a remarkable sexual dimorphism concerning the contents of porphyrins and melatonin. Mainly for this reason, the hamster HG has been used as a model to compare, under normal conditions, the physiological oxidative stress between females (strong) and males (moderate). Androgens are responsible for the sexual dimorphism in hamster and they are known to control the HG secretory activity in different species. Furthermore, HG is a target of pituitary, pineal and thyroid hormones. This review offers a comparative panorama of the endocrine activity of the HG as well as the hormonal control of its secretory activity, with a particular emphasis on the sex dimorphic aspects of the hamster HG.

Ultrastructure of plasma cells in harderian gland of laying hens.

Ultrastructure of plasma cells in Harderian gland was investigated using the transmission electron microscopy. For this research, we examined the glands of 32 laying hens collected at 1, 7, 20 and 40 days and 4, 6, 8 and 12 months of the birds' ages. The research showed that the stroma of the gland contains a large number of lymphocytes and plasma cells. Most of the plasma cells are mature, but morphologically do not show productive activity. Only some individual plasma cells, situated under the secretory epithelium of primary and secondary ducts, have extremely dilated cisternae of rough endoplasmic reticulum which contain moderately dense, granular material. The morphology of these cells indicates that they are in active stage of immunoglobulin production. Also, we identified plasma cells with two types of Russell bodies. One type of these bodies was small, round or oval, while the other had irregular, angular shape. It was noted that one plasma cell never contains both type of Russell bodies at the same time. These cells were often affected by apoptosis. Among them, in deeper part of the stroma, were situated the small plasmablast cells.

Secretory glands and microvascular systems imaged in aqueous solution by atmospheric scanning electron microscopy (ASEM).

Exocrine glands, e.g., salivary and pancreatic glands, play an important role in digestive enzyme secretion, while endocrine glands, e.g., pancreatic islets, secrete hormones that regulate blood glucose levels. The dysfunction of these secretory organs immediately leads to various diseases, such as diabetes or Sjögren's syndrome, by poorly understood mechanisms. Gland-related diseases have been studied by optical microscopy (OM), and at higher resolution by transmission electron microscopy (TEM) of Epon embedded samples, which necessitates hydrophobic sample pretreatment. Here, we report the direct observation of tissue in aqueous solution by atmospheric scanning electron microscopy (ASEM). Salivary glands, lacrimal glands, and pancreas were fixed, sectioned into slabs, stained with phosphotungstic acid (PTA), and inspected in radical scavenger d-glucose solution from below by an inverted scanning electron microscopy (SEM), guided by optical microscopy from above to target the tissue substructures. A 2- to 3-µm specimen thickness was visualized by the SEM. In secretory cells, cytoplasmic vesicles and other organelles were clearly imaged at high resolution, and the former could be classified according to the degree of PTA staining. In islets of Langerhans, the microvascular system used as an outlet by the secretory cells was also clearly observed. Microvascular system is also critically involved in the onset of diabetic complications and was clearly visible in subcutaneous tissue imaged by ASEM. The results suggest the use of in-solution ASEM for histology and to study vesicle secretion systems. Further, the high-throughput of ASEM makes it a potential tool for the diagnosis of exocrine and endocrine-related diseases.

Histological, histochemical and ultrastructural studies on Harderian and lacrimal glands of the Capercaillie (Tetrao urogallus major L.).

This study describes the macroscopic anatomy and the microscopic and ultrastructural features of the Harderian gland and lacrimal gland of the Capercaillies. It was conducted both on adult male and female Capercaillies. Tissue sections were stained with hematoxylin and eosin, azan trichrome, modified Mallory's trichrome, methyl green-pyronin Y, periodic acid-Schiff, alcian blue pH 2.5, aldehyde fuchsin and Hale's dialysed iron. The morphometric study of the Harderian and lacrimal glands indicated that they are both larger in male than in female Capercaillies. The histological analysis showed that the HG has a multilobar tubulo-alveolar structure with numerous lymphocytes and plasma cells. The LG has a multilobar tubulo-acinar structure without lymphocytes and plasma cells. The periodic acid-Schiff staining and alcian blue pH 2.5 staining demonstrated a mild positive reaction in the epithelial cells of the Harderian gland and weak positive reaction in the lacrimal gland. The HDI staining detected the presence of carboxylated acid mucopolysaccharides in the Harderian and lacrimal glands. Transmission electron microscopy revealed the presence of two types of secretory vesicles in the cytoplasm of both studied glands. It also showed that lipid droplets and glycogen granules were more abundant in the Harderian gland than in the lacrimal gland of this species.

Structure of the secretory cells of male and female adult guinea pigs Harderian gland.

The main objective of this study was to investigate the structure of the Harderian gland (HG) in male and female guinea pigs. A total number of sixteen animals of 4 months age were divided according to sex into two groups; eight animals each. Unfixed glands were weighed and their length and width were measured. Specimens from fixed glands were processed and examined using light, transmission electron microscopy and immunohistochemistry for the detection of the presence of chromogranin A (CgA). The gland consisted of a well-developed duct system which included both intra and extra parenchymal ducts and secretory end pieces lined by many types of cells of variable morphological features and modes of secretion. However, the holocrine mode of secretion was rare as mitotic figures were occasionally present. The interstitial cells included fibroblasts and immune cells (mast cells, lymphocyte, plasma cells and macrophages). The secretion produced by the gland included lipid, protein, neutral mucin and CgA which may be a newly identified constituent of biologically potent proteins stored in the cells of the guinea pig HG. Neutral mucin and CgA may function in photoprotection. The gland revealed sexual dimorphism in mast cells and blood capillaries number and chromogranin secretory activity.

Analysis of constant tissue remodeling in Syrian hamster Harderian gland: intra-tubular and inter-tubular syncytial masses.

The Syrian hamster Harderian gland (HG) has a marked sexual dimorphism and exhibits an extraordinary rate of porphyrinogenesis. The physiological oxidative stress, derived from constant porphyrin production, is so high that the HG needs additional survival autophagic mechanisms to fight against this chronic exposure, provoking the triggering of a holocrine secretion in female glands that forms two types of secretory masses: intra-tubular-syncytial and inter-tubular-syncytial masses. The aim of this work was to study the development of this inter-tubular holocrine secretion. To approach this task, we have considered that the steps developed during the formation of the so-called invasive masses consist of the growth of epithelial cells, cell detachment from the basal lamina and invasion of surrounding tissues. The presence of these masses, particularly in the female HG, are closely linked to sexual dimorphism in redox balance and to alterations in the expression of certain factors such as cytokeratins, P-cadherin, matrix metalloproteinases, cathepsin H, proliferating cell nuclear antigen, p53, CD-31 and vascular endothelial growth factor, which seem to be involved in tissue remodeling. The results document unusual mechanisms of secretion in Syrian hamster HG: an extraordinary system of massive secretion through the conjunctive tissue, disrupting the branched structure of the gland.

Light and electron microscopic studies of the Gerbillus tarabuli (Thomas, 1902) Harderian gland.

The purpose of this investigation was to study the morphological aspects of the Harderian gland in Gerbillus tarabuli. Tissues were obtained from both male and female adult Gerbillus tarabuli and processed for light and electron microscopy. The Harderian gland in gerbil is large and well developed, covered by a thin capsule, from which thin septae extend, subdividing the gland into lobes and lobules. The endpieces of the gland are tubuloalveolar, which produce a secretion of lipid character. The glandular epithelium is pseudostratified with two types of secretory cells, the type C cells are columnar in shape with large lipid vacuoles, and type P cells pyramidal and serous, they are basally located with no luminal aspect. The epithelium possesses well-developed myoepithelial cells. The wide lumina are filled with lipid vacuoles, cellular debris, and porphyrins. The Harderian gland of the gerbil has no morphologically distinct duct system; a single extraglandular excretory duct is detected. Electron microscopic examination revealed that type C cells contain large electron-light lipid vacuoles, a well and extensive reticulum endoplasmic and a large number of mitochondria. The pyramidal cells are characterized by a small number of PAS-positive granules at the basal region; these cells exhibit one or two round nuclei, many electron-dense granules, crystalloid bodies, abundant mitochondria and many ribosomes in their cytoplasm. The three mechanism of secretion are seen in the Harderian gland of Gerbillus tarabuli. In its overall characteristics, the Harderian gland of Gerbillus tarabuli conforms to the general pattern observed in rodents. However, further research will be needed to correlate the presence of cytoplasmic slashes, crystalloids bodies and glycoproteins in epithelial cells with the biology of these animals and to their functional significance.

Is it or isn't it? A reexamination of the anterior orbital glands of the fat-tailed Dunnart Sminthopsis Crassicaudata (Dasyuridae: Marsupiala) and a reevaluation of definitions for the Harderian gland.

The anterior orbital glands of tetrapods, which include the Harderian and nictitans glands, can usually be differentiated either anatomically (nictitans gland is more anterior) or histochemically (Harderian gland secretes lipids). However, conflicting statements exist in the literature about the presence and identity of these glands. Two previous studies on Sminthopsis crassicaudata (Dasyuridae: Marsupiala) either failed to note any anterior ocular glands or used no histochemical analyses. This study reexamined the structure of the anterior orbital glands of S. crassicaudata. Histological, histochemical, and ultrastructural examination revealed three glandular units: two of which are located superficially in the nictitating membrane, the third lying deeper in the connective tissue. The ducts of these three glandular units all open onto the corneal aspect of the nictitating membrane. These cells contain mainly serous granules with sparse intracellular lipid droplets. The nomenclature of these structures depends upon the definition used. According to the anatomical definition, S. crassicaudata has two glands: anteriorly the nictitans and posteriorly the Harderian gland. In contrast, if the histochemical definition is used, there is only one gland, but its precise identity cannot be confirmed until the role of the lipid droplets is established. Moreover, the histochemical definition poses additional problems with respect to the mechanism of secretion, multiple secretions, and glandular plasticity. Finally, the unitary definition identifies one deeply subdivided gland with an anterior and a posterior lobe in S. crassicaudata. This last definition is broad enough to accommodate a wide level of anatomical variation in the anterior ocular glands of tetrapods.

Histomorphology of the Harderian gland in the Angora rabbit.

This study was aimed to demonstrate the morphological and histochemical properties of the Harderian gland in the Angora rabbit. Ten healthy adult Angora rabbits obtained from private breeders constituted the material of the study. The Harderian gland, which is composed of the pink and white lobes, consists of cells that produce a secretion of lipid character. The pink lobe contained type I cells with large lipid vacuoles. Cells with small lipid vacuoles (type II) were found in the white lobe. Type III cells containing both large and small lipid vacuoles were not observed. While type I cells reacted strongly to staining with Oil red O, type II cells reacted weakly to this stain. The number of plasma cells was greater in the white lobe when compared to the pink lobe. The apical granules within the epithelial cells lining the intralobular and inter-lobular excretory ducts of the gland were positive for periodic acid-Schiff (PAS), periodic acid-Schiff/alcian blue (PAS/AB), alcian blue (AB) and performic acid/alcian blue (PA/AB). Electron microscopic examination revealed that type I cells contain large electron-light lipid vacuoles and an eccentric heterochromatic nucleus, due to the presence of these vacuoles. The cells, which were connected by tight junctions, possessed apically located microfolds. The nucleus of type II cells was situated basally and had an oval shape. Type II cells had apical microvilli-like cytoplasmic protrusions, longer than those of type I cells. Oval shaped myoepithelial cells were observed between the glandular epithelial cells and their basal lamina. The epithelium lining the excretory ducts of the gland contained two types of granules, which were dark and lightly coloured. Histochemical and ultrastructural examinations revealed no difference in the structure of the Harderian gland between female and male Angora rabbits.

Underdeveloped extraocular muscles in the naked mole-rat (Heterocephalus glaber).

The extraocular muscles (EOM), the effector arm of the ocular motor system, have a unique embryological origin and phenotype. The naked mole-rat (NMR) is a subterranean rodent with an underdeveloped visual system. It has not been established if their ocular motor system is also less developed. The NMR is an ideal model to examine the potential codependence of oculomotor and visual system development and evolution. Our goal was to compare the structural features of NMR EOMs to those of the mouse, a similar sized rodent with a fully developed visual system. Perfusion-fixed whole orbits and EOMs were dissected from adult NMR and C57BL mice and examined by light and electron microscopy. NMR orbital anatomy showed smaller EOMs in roughly the same distribution around the eye as in mouse and surrounded by a very small Harderian gland. The NMR EOMs did not appear to have the two-layer fiber distribution seen in mouse EOMs; fibers were also significantly smaller (112.3 +/- 46.2 vs. 550.7 +/- 226 sq microm in mouse EOMs, *P < 0.05). Myofibrillar density was less in NMR EOMs, and triad and other membranous structures were rudimentary. Finally, mitochondrial volume density was significantly less in NMR EOMs than in mouse EOM (4.5% +/- 1.9 vs. 21.2% +/- 11.6, respectively, *P < 0.05). These results demonstrate that NMR EOMs are smaller and less organized than those in the mouse. The "simpler" EOM organization and structure in NMR may be explained by the poor visual ability of these rodents, initially demonstrated by their primitive visual system.

Scanning electron microscopy of the orbital Harderian gland in the male Atlantic bottlenose dolphin (Tursiops truncatus).

The ultrastructure of the Harderian gland of Atlantic bottlenose dolphin (Tursiops truncatus) was examined by scanning electron microscopy (SEM). We found the following surface features: the typical round appearance of the ascinar glandular unit with a finely granular surface, a thin cortex and immediately below two types of cells: type I cells (characterized by small lipid vacuoles) and type II cells (characterized by large lipid vacuoles). It has been suggested that different cells forms represent a single cell type in varying activity states. Additionally, a coalescent tubular complex, a small balloon-like structures and large globular structures were observed. These structures may be reservoirs of secretion products.

D-Aspartate affects secretory activity in rat Harderian gland: molecular mechanism and functional significance.

In this paper, the role of D-aspartate in the rat Harderian gland (HG) was investigated by histochemical, ultrastructural, and biochemical analyses. In this gland, substantial amounts of endogenous D-Asp were detected, along with aspartate racemases that convert D-Asp to L-Asp and vice versa. We found that the gland was capable of uptaking and accumulating exogenously administered D-Asp. D-Asp acute treatment markedly increased lipid and porphyrin secretion and induced a powerful hyperaemia in inter-acinar interstitial tissue. Since D-Asp is known to be recognized by NMDA receptors, the expression of such receptors in rat HG led us to the hypothesis that D-Asp acute treatment induced the activation of the extracellular signal-regulated protein kinase (ERK) and nitric oxide synthase (NOS) pathways mediated by NMDA. Interestingly, as a result of enhanced oxidative stress due to increased porphyrin secretion, the revealed activation of the stress-activated protein kinase/c-jun N-terminal kinase (SAPK/JNK) pro-apoptotic pathway was probably triggered by the gland itself to preserve its cellular integrity.

Morphological and biochemical changes in the Harderian gland of hypothyroid rats.

The secretory activity of the Harderian gland (HG) is influenced by both exogenous (such as light and temperature) and endogenous (such as prolactin, thyroid hormones and steroid hormones) factors, which vary among species. In the present study, the effects of hypothyroidism on the rat HG were examined at morphological and biochemical levels. The decrease in cytoplasmic lipoproteic vacuoles and the increase in mucosubstance secretion in the acinar lumina were the most notable histological effects elicited by hypothyroidism. The release of all granules with nuclei and cellular debris suggested the occurrence of holocrine secretion. Electron microscopy revealed in the glandular cells of hypothyroid rat an increased condensation of chromatin in the nuclei, mitochondria with decreased cristae and vacuolisation, decreased glycogen granules, autophagic vacuoles, and lipofuscins in the cytoplasm. TUNEL reaction indicated DNA fragmentation in hypothyroid HG, indicative of an underlying apoptotic process. Translocation of cytochrome c from mitochondria to cytosol strongly supported this hypothesis. In conclusion, these findings indicate that thyroid hormones play a pivotal role in preserving the structural integrity of the rat HG and, hence, its secretory activity.

The orbital Harderian gland of the male atlantic bottlenose dolphin (Tursiops truncatus): a morphological study.

The ultrastructure of the Atlantic Bottlenose dolphin Harderian gland (HG) has been described but some questions remain unanswered. The purpose of this work was to define the gland's structure, ultrastructure and the differences between cells (types I and II) of the male dolphin using optic, fluorescence and electron transmission microscopy. Three different cells were observed under optic and fluorescence microscopic examination, while only two cell types (types I and II) were distinguished by electron transmission microscopy. Type I (oval nuclear envelope) exhibited three different cell populations and type II (indented nuclear envelope) exhibited two different cell populations. Although, we observed both types of vesicles in both types of cells they differed, principally, in quantity. The glands also possessed prominent duct systems, with three orders of complexity. The dolphin orbital HG appears to function as a mixed heterologous gland with two types of cells that exhibit both types of vesicles and other distinguishable differences.

Expression of different classes of immunoglobulin in intraepithelial plasma cells of the Harderian gland of domestic ducks Anas platyrhynchos.

The Harderian gland of chickens contains numerous plasma cells and is considered as a peripheral lymphoid organ. Data about this gland in other avian species are scarce or inexistent. Considering that ducks show some unique characteristics regarding the immune system, which are important in evolutionary context, and that unusual location of plasma cells into the epithelium was recently described in primitive avian species, here we investigated the occurrence and characterized intraepithelial plasma cells in the Harderian gland of ducks, according to the immunoglobulin produced. Numerous intraepithelial plasma cells were found confined to the Harderian gland ducts. Plasma cells were also found in the ducts lamina propria. IgM-positive cells were the most abundant into the epithelium. In contrast, IgY- or IgA-positive cells were predominant in the lamina propria. The constancy of intraepithelial plasma cells in all specimens examined indicates that they may be essential mediator for an effective immunesurvaillance of the ocular mucosa.

Survival mechanisms in a physiological oxidative stress model.

The Syrian hamster Harderian gland has as the remarkable feature of an extraordinary rate of porphyrin production, even higher than the liver. The low activity of the last enzyme of the route gives rise to the accumulation of the uncomplex porphyrins in the female glands. Moreover, due to the localization of the Harderian gland, porphyrins exposed to light produce reactive oxygen species and, thus, the gland presents a physiological oxidative stress, with a great number of sings of degeneration, but without compromising the gland integrity. The appearance of abnormal features in this gland was largely described in the past, but the significance is interpreted for the first time in this study. We have found that autophagic processes are the first result of an elevated porphyrin metabolism, as it is observed in both sexes. This mechanism is considered, in this case, as a constant renovation system that allows the normal gland activity to be sustained. Furthermore, there is a second procedure, invasive processes toward connective tissue, which even occasionally reach blood vessels with intravasation of damaged gland components into the bloodstream. This effect is a consequence of a strong oxidative stress environment that is mainly observed in the female gland, resembling to tumoral progression. Both mechanisms, autophagy and invasive processes, have to be implied in the maintenance of the gland integrity.

Anatomy, histology, histochemistry and fine structure of the Harderian gland in the South American armadillo Chaetophractus villosus (Xenarthra, Mammalia).

The anatomical, histological, histochemical and ultrastructural characteristics of the Harderian gland of the armadillo Chaetophractus villosus were described. The gland is the largest structure in the bony orbit. It is situated in the anteroventral region of the orbit. Obvious structural differences are not observed between males and females. The gland is compound-branched tubulo-alveolar, being characterized by a single layer of columnar cells surrounded by myoepithelial cells. It possesses a single excretory duct opened into the inner canthus. All glandular cells show yellow-green autofluorescence and additionally some glandular lumen may contain dense autofluorescent solid accretions. There are two peculiar and outstanding cytoplasmic features. One is represented by the smooth endoplasmic reticulum (SER), forming a closely woven meshwork. The other one is represented by "membranous bodies" apparently derived from the SER, RER and cytoskeleton with a "Star of David" configuration situated in the supranuclear region. Three types of vesicles are detected in the cytoplasm. Histochemical staining methods reveal lipids, proteins, neutral and acidic containing glycoconjugates in secretory vesicles. The mechanism of secretion appears either merocrine or apocrine. The epithelium of the intra- and inter-lobular excretory ducts suggests secretory activity. Tubulo-acinar glands similar to those seen in the lacrimal gland and nictitans glands are found related to the intralobular and main excretory ducts. The capillary network is characterized by fenestrated endothelium. The stroma possesses unmyelinated axons and plasma cells. The normal secretion of the secretory endpieces, particularly lipids, proteins and glycoconjugates, is complemented by mucous and serous secretions released by ductal cells and glands associated to the ducts.

D-aspartate modulates transcriptional activity in Harderian gland of frog, Rana esculenta: Morphological and molecular evidence.

In the green frog, Rana esculenta, a substantial amount of D-aspartate (D-Asp) is found endogenously within the Harderian gland (HG) following its synthesis from L-aspartate (L-Asp) by an aspartate racemase. The frog HG is an orbital seromucoid gland that displays seasonal changes in secretory activity. Our in vivo experiments, consisting of i.p. injection of 2.0 mumol/g b.w. D-Asp in frogs collected during two periods of differing glandular activity (high or medium-low secretory activity), revealed that HG can to take up and accumulate D-Asp and that this amino acid may modulate the exocrine secretion through a kinase pathway. At a time when the gland shows relatively low secretory activity, i.p. administration of D-Asp rapidly induced activation of ERK1 and an increase in cells active in RNA synthesis. This increase in transcriptional activity was followed by a significant increase in mucous secretion. By contrast, administration of exogenous D-Asp when HG was showing high activity rapidly induced inhibition of both ERK1 and transcriptional activity. Since D-Asp is known to be recognized by receptors for N-methyl-D-aspartic acid (NMDA), it is possible that in the HG, D-Asp mediated NMDA activation may enhance the kinase pathway. The above activation of opposing stimulatory and inhibitory processes could reflect different levels of NMDA-receptor activity, which could vary as a function of the level of gland activity. This study provides the first evidence of a role for this excitatory amino acid in exocrine secretion. The effects of D-Asp in HG appear to be specific since they were not seen in frogs treated with other D- or L-amino acids with known excitatory effects on neurosecretion.

The Harderian gland of the Mexican volcano mouse Neotomodon alstoni alstoni (Merriam 1898): a morphological and biochemical approach.

The Harderian glands of rodents are large intraorbital exocrine glands with histologic organization that varies among mammalian species. Here we describe some ultrastructural and biochemical features of the Harderian gland in the Mexican volcano mouse Neotomodon alstoni alstoni, a species of restricted habitat. The Harderian glands from male and female adult mice were dissected, processed and embedded in Epon 812 for light and electron microscopy studies. Porphyrin and total lipids were biochemically determined. The macroscopic appearance of the Harderian gland is similar in the male and female. The gland is a bilobulate structure, situated in the orbit towards the posterior side of the eyeball, of whitish color and is surrounded by a connective tissue capsule. The male gland is slightly heavier (127 mg) than that of the female (113 mg). The Harderian gland shows a tubulo-alveolar organization and is composed exclusively of one type of secretory cells. No branched duct system within the gland was found. Adrenergic nerves endings and mast cell were observed in the interstices of the alveoli. Male and female glands produce similar levels of porphyrins. Triglyceride levels were significantly higher (P < 0.05) in the female compared to the male. Abundance of lipids could induce corneal lubrication of the Harderian gland which may confer a protective and adaptative function to the volcano mouse in its natural habitat during the dry and cold seasons.

The fine structure of the Harderian gland in the ostrich (Struthio camelus).

The Harderian gland of the ostrich (Struthio camelus) is a tubuloalveolar gland containing holocrine secreting epithelial cells. The gland epithelium is composed of two different cell types, which can be classified as type I and type II. These cells contain dense secretory vesicles in their cytoplasm and they are connected laterally with desmosomes. At the basal site of these cells, myoepithelial cells are present. Plasma cells are observed in the subepithelial region of the gland. In the interlobular trabeculae, forming the gland stroma, fibroblasts, blood vessels and nerve fibres are included. Another important finding in the ostrich Harderian gland is the presence of homogeneous material.