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The very high rate of protein synthesis in acinar cells generates a constant low level of ER stress. Science, Technology and Medicine open access publisher.Publish, read and share novel research. The Endocrine Glands in the Dog: From the Cell to HormoneHelena Vala1, 2, Joao Rodrigo Mesquita2, Fernando Esteves2, Carla Santos2, Rita Cruz2, Cristina Mega2 and Carmen Nobrega2[1] Center for Studies in Education, and Health Technologies. Pancreas resection - A portion of a person's pancreas (or in some cases the entire pancreas) is surgically removed usually because they have a cancerous tumor. Pancreas divisum: Pancreas divisum is a common congenital anomaly (an anomaly that is present at birth) of the pancreatic duct(s). The cardiac sphincter between the esophagus and the stomach: If this must be removed then the patient will have reflux from the stomach into the esophagus, especially if lying down too soon after eating a meal.
The pyloric sphincter between the stomach and the duodenum: If this must be removed then the stomach will empty quickly after eating, reducing both digestion and absorption of food in the stomach and potentially causing dumping syndrome, also called early gastric emptying. Pre-surgery anatomy (top panel) and pancreaduodenectomy either sparing the pylorus (second panel) or with resection of the pylorus (third panel). The sphincter of Oddi, or otherwise called the sphincter of the hepato-pancreatic ampulla, is located at the second part of the duodenum. The migrating motor complex, or the MMC for short, is smooth muscle contractions that are also termed myoelectrical motor complex.
Another function of the sphincter of Oddi is to prevent entry of contents of the duodenum into the bile duct or pancreatic duct.
Although the sphincter of Oddi is just one part of the extra-hepatic biliary tree, its functions are very vital to maintain order inside the human body. During surgical removal of the gallbladder or even in gallstone removal, iatrogenic trauma may be caused to the sphincter of Oddi. Under normal conditions, this ER stress is readily compensated for by the UPR and EOR, which reduce the rate of protein synthesis while increasing production of molecules that facilitate cell recovery and repair.
ThyroidCanine thyroid gland is located at cervical ventral region, lateral and ventral to the 5th-8th tracheal rings (Figure 7), being composed by two separate lobes, occasionally connected by an isthmus [17, 18].
Adrenal glandsThe adrenal glands are paired organs located against the roof of the abdomen near the thoracolumbar junction, in a position immediately prior to the kidneys and close to their cranial poles (Figure 16). Clinical significance Endocrine diseases, associated to altered functions of endocrine glands, are frequently seen in veterinary practice.
Molecular and genetic study of the role of hormones receptors, and enzymes in regulation of reproduction, lipid metabolism, and other human physiological functions.
Follicle-Stimulating Hormone and Luteinizing Hormone Mediate the Androgenic Pathway in Leydig Cells of an Evolutionary Advanced Teleost. G proteins and autocrine signaling differentially regulate gonadotropin subunit expression in the pituitary gonadotrope. Thyroxine-induced expression of pyroglutamyl peptidase II and inhibition of TSH release precedes suppression of TRH mRNA and requires type 2 deiodinase.
Secretory cells of the supraoptic nucleus have central as well as neurohypophysial projections. Physiology of the pancreatic ?-cell and glucagon secretion: role in glucose homeostasis and diabetes. After surgery patients may need pancreatic enzyme supplements and insulin depending on how much their body can produce with a reduced pancreas. The end of the remaining sigmoid colon is brought out through the surface of the abdomen in a permanent colostomy. As the name implies, it houses the ampulla of Vater, which in turn surrounds the bile duct and pancreatic duct. Its tonic contraction creates a basal pressure higher compared to the pressure in the common bile duct and duodenum.
The substance cholecystokinin relaxes the sphincter of Oddi along with the stimulation of pancreatic enzyme secretion and contraction of the gallbladder. It is always important to go back to the anatomy of the sphincter in order to understand how it is able to perform all these functions at the same time. If the sphincter of Oddi fails to perform even one of its functions, it would have devastating effects to digestion and to the quality of life of the person. This would cause inflammation to the sphincter which in its process of repair would eventually cause narrowing.
Agrarian School of Viseu, Polytechnic Institute of Viseu, Viseu, Portugal[2] Agrarian Superior School of Viseu, Polytechnic Institute of Viseu. In the dog there are four parathyroid glands, one external and one internal per each thyroid gland [17, 19, 20].External parathyroid glands are capsulated and may be found in varied positions, according to the species, which means that they can be placed between a cranial location to the thyroid gland and the entrance of the chest.
The endocrine glands described may be targeted by several conditions summarized in table 7. Two histologic forms of primary hypothyroidism predominate in the dog: lymphocytic thyroiditis and idiopathic atrophy. After two hours or more in the stomach, the partly digested food moves into the beginning of the duodenum. One of its functions is to produce enzymes that are important for the digestion of food in the intestine. Windham's tutorial Abdominal Anatomy for GIST in our website's Ask the Professional section.
The key in order to understand the function of the sphincter of Oddi is to discuss first how it looks like. It is produced by the mucosal cells that lines the upper small intestine once they detect presence of fats and amino acids. Studies on the physiology of the sphincter of Oddi show that the interplay between the small component sphincters and the longitudinal muscle fibers.
Because the sphincter of Oddi relaxes after a meal when it is highly stimulated by cholecystokinin, pain from its dysfunction is mostly felt after eating and is called postprandial pain. Other effects are also paradoxical response of the sphincter of Oddi to cholecystokinin resulting to dyskinesia or non-relaxation. Excessive acinar cell apoptosis can lead to necrosis, a severe systemic inflammatory response, acute respiratory distress, and life-threatening multiorgan failure.
When the food reaches the duodenum, the pancreas releases its digestive juices which flow down the pancreatic duct and mix with the food.
The digestive enzymes, in the form of digestive juice, drain from the pancreas via the pancreatic duct into the duodenum (the upper portion of the small intestine) where they aid in digesting food. This is to ensure that contents of the common bile duct and pancreatic duct do not enter the duodenum. It has three phases, the third phase of which migrates from the antrum of the stomach down to the ileum of the small intestine. In between contractions, the sphincter of Oddi allows flow of bile and digestive enzymes into the duodenum.
Because of its function as a pump for bile and pancreatic secretions, its failure to do so frequently causes pancreatitis because the digestive enzymes of the pancreas are not secreted and are instead directed on the organ causing inflammation. Altered Ca2+ release from the ER, and ROS, are thought to activate NF-κB, a key regulator of inflammatory gene expression. IntroductionThe animal body represents one of the more complex and perfect systems of nature. They may be absent in some species or may be found included within the thyroid or close to them.
The slender right lobe runs within the mesoduodenum and the thicker but shorter left lobe extends over the caudal surface of the stomach towards the spleen, within the greater omentum (Figure 13) [4, 17].
The cortex is yellowish and radially striated, while the medulla is more uniform and darker [17].
Doberman Pinschers and Golden Retrievers are the two breeds most frequently described [42].


Take note that the pancreas secretes digestive enzymes that may be destructive to the mucosa of the gut when uncontrolled by the sphincter of Oddi. The longitudinal fibers of the sphincter of Oddi previously mentioned function as promoters of the flow of bile and digestive enzymes into the duodenum.
The sphincter of Oddi also responds to motilin, which controls the motility of the digestive tract.
This mechanism, which is termed as trans-sphincteric flow, shows that the sphincter performs its resistor and pump functions simultaneously. Despite its complexity and its functionality, which is incredibly effective, the control of its basic functions is performed by only two systems: the nervous system and endocrine system. Bar = 100 µm.The parenchyma of the pars distalis is composed by chromophilic cells, acidophilic or basophilic, arranged in cords, nests or follicles (Figure 4) and by small, round endocrine cells, without evident granules - chromophobe cells which means that they have low tinctorial affinity, also known as principal cells, reserve cells or C cells [4].
Bar = 25 µm.The follicle is, therefore, the structural and secretory unit of the thyroid gland, whose center is filled with a colloidal secretion (Figure 9), comprising the hormones triiodothyronine (T3) and thyroxine (T4) [4]. In the dog, internal parathyroid glands are within the thyroid capsule, in the caudal and medial aspects of the thyroid [19, 20].The parenchyma of parathyroid glands is composed of secretory cells, named chief or principal cells, arranged in cords, clusters, chains or rosettes (Figure 11, Figure 12). Both regions, cortex and medulla, correspond to areas specialized in producing different hormones [4, 32, 34]:The cortex consists of polyhedral secretory cells, arranged into two layered thick cords, which originate radially from the medullar zone [4, 33]. Clinical signs associated with hypothyroidism are varied, and behavioural changes are associated with a reduced metabolic rate, dermatological signs, cardiovascular and neuromuscular abnormalities. In more than 90% of the embryos, the dorsal and the ventral ducts will fuse to form one main pancreatic duct.
Longitudinal muscle fibers or fasciculi longitudinales are found in between the bile and pancreatic ducts.
The sphincter’s contractions increase in frequency and amplitude shortly before and during the intense contractions of the duodenum. Trans-sphinteric flow is diminished by the action of motilin, with which the sphincter of Oddi also responds. The nervous system is associated with electrical and chemical signals that are transmitted at high speed, resulting in rapid organic activities. Chromophilic cells secrete somatotropic hormone (also known as somatotropin - STH or growth hormone - GH), prolactin (PRL), adrenocorticotropic hormone (ACTH), follicle-stimulating hormone (FSH), luteinizing hormone (LH) (which is also called interstitial cell-stimulating hormone or ICSH in males), thyrotropin or thyroid-stimulating hormone (TSH), melanocyte-stimulating hormone (MSH) and lipotropines [4].
In between the follicles, in a parafollicular position, cells with a pale cytoplasm, and a basal nucleus can be found - parafollicular cells or C cells which produce calcitonin [4, 16, 18].
In some species, such as cattle and humans, it is also composed by oxyphilic cells, organized in small clusters which have yet no known functions [4].
The later is produced in the islets of Langerhans, which are randomly scattered in the exocrine parenchyma (Figure 14) [10, 17, 24]. The main pancreatic duct will join the common bile duct (the duct that drains bile from the gallbladder and the liver) to form a common bile and pancreatic duct which drains into the duodenum through the major papilla.
The endocrine system acts through the synthesis and release of chemical messengers and is responsible for several functions of the organism, in a slower, but more durable way.Endocrinology is the science that studies the internal secretions produced by endocrine glands. Somatotropin promotes epiphyseal growth [6] and protein production, whereas prolactin leads to mammary gland development and milk secretion. In dogs, this type of cells were only found in senile animals [21] and were also described in canine parathyroid adenoma [22]. Pancreatic islets are aggregates of endocrine cells required for blood glucose control and diabetes prevention after birth, consisting of polygonal shaped cells, with pale eosinophilic cytoplasm and coarse heterochromatin. The outer zone is the zona glomerulosa, a thin region that in dogs and cats is composed by cells arranged in an arched or arcuate pattern, therefore also named as zona arcuata (Figure 17) [32, 35, 36].The zona glomerulosa produces mineralocorticoids, particularly the steroid hormone aldosterone [32, 37], which is responsible for increasing sodium retention and water reabsortion in the distal tubules of the kidneys and sodium reabsorption in intestine. Endocrine glands are distributed throughout the body and secrete chemical messengers – hormones, in response to an internal or external stimulus. ACTH acts on the adrenal gland cortex, resulting in an increased release of adrenocortical hormones [7]. Cells are arranged in cords or clusters separated by sinusoids and are subdivided in the following subtypes (Figure 15) [4, 24, 25]:? cells, located peripherally and representing 10-20% of islet cells, secrete glucagon, a hyperglycaemic hormone, as well as cholecystokinin, gastric inhibitory protein and ACTH-endorphin [4, 10, 24, 26].
Aldosterone also maintains blood concentration and stimulates potassium excretion by the kidneys, thereby, indirectly regulating extracellular fluid volume. Seborrhoea, bilateral and symmetrical alopecia, and pyoderma are also common dermatological symptoms. Failure of the ventral and the dorsal pancreatic ducts to fuse is called pancreas divisum (because the pancreas is drained by two ducts). These hormones are released directly into the bloodstream – endocrine mechanism, in contrast to exocrine glands, which use a ductal system to release their secretions in locations that lead, ultimately, to the exterior of the body – exocrine mechanism. ? cells, located in the centre of the islets, are more abundant, representing 60-80% of the islets of Langerhans, and secrete insulin, a hypoglycaemic hormone [4, 10, 24].
A decrease of mineralocorticoids, by loss of this zone or its functional ability, gives rise to water outlet of the blood to the tissues and may result in death, due to retention of high levels of potassium with excess loss of sodium, chloride and water [32].
These signs can be accompanied by bradycardia, low voltage ECG complexes and weakness [40, 41, 43, 44].Diabetes mellitusDiabetes mellitus is the result of any situation that affects insulin production, insulin transport or the sensitivity of target tissues to insulin [45].
In pancreas divisum, the ventral duct drains into the major papilla, while the dorsal duct drains into a separate minor papilla. Hormones are transported through the bloodstream to target organs, where they will exert a physiological control, even in low concentrations, coordinating a multiplicity of organic functions and maintaining homeostasis. Many factors are known to contribute to the development of diabetes and its complications [46, 47].
The main endocrine glands in the animal body include pituitary gland, thyroid, parathyroid, pancreas, adrenal (Figure 1), and gonads (ovaries and testes).
This hormone is also capable of reducing intestinal motility and secretion of digestive juices. These include genetics, diet, sedentary lifestyle, perinatal factors, age, obesity and inflammatory causes [47, 48]. A small number of patients with pancreas divisum will experience repeated episodes of pancreatitis.
These cells represent about 5-40% of the islets [10, 24].Besides the above-mentioned cell types, the endocrine pancreas also contain some minor types of cells, that represent around 5% of the total pancreatic cells. Canine diabetes mellitus is generally classified as insulin dependent or non-insulin dependent based on the need for insulin treatment [40, 48]. Pancreatitis is an inflammation of the pancreas that can cause severe abdominal pain and more severe complications.
Gonadotropes or gonadotropic cells which secrete gonadotropins LH and FSH (basophilic) [13].
This hormone, together with calcitonin and vitamin D, is involved in the regulation of calcium homeostasis. Bar = 50 µm.The zona fasciculata is the middle and thickest zone (corresponding to more than 70% of the cortex) and is composed of parallel columns of secretory cells, one to two cells thick, separated by prominent capillaries.
Most diabetic dogs are thought to have a disease like human type I diabetes mellitus and are insulin dependent [40, 49, 50]. Hypothalamic-pituitary axisThe hypothalamus, located at the base of the ventral diencephalon is limited rostrally by the optic chiasm, caudally by the mammillary processes, laterally by the temporal lobes and dorsally by the thalamus.
It is released in response to low blood calcium level [23] due to it’s ability to enhance the mobilization of calcium ions from the small intestine and from bone resorption, and also by increasing the simultaneous absorption of calcium and excretion of phosphate from distal convoluted tubules of kidneys [10] (Table 4). The cells are cuboidal or polyhedral, containing vesicular nucleus, frequently binucleated) and foamy cytoplasm (intracellular lipid droplets) (Figure 18). Unlike human type I diabetes mellitus, which occurs mainly in young human patients, in dogs it is more likely to occur later in life [40, 45, 48]. The chromophobe cells are also arranged in clusters or cords and their low tinctorial affinity probably indicates that these cells may correspond to depleted cells of any of the above types described or a state of partial degranulation.
Dogs do not appear to progress from obesity-induced insulin resistance to type 2 diabetes mellitus, probably because pancreatic beta cells in dogs are either not sensitive to toxicity due mild hyperglycemia or lack other components of the pathophysiology of beta cell failure in type 2 diabetes mellitus [51]. One theory is that the minor papilla is too narrow to adequately drain the digestive juices in the dorsal duct.


Although not considered a real endocrine gland, the hypothalamus coordinates all pituitary activity by the release of a number of peptides and amines that control the secretion of hormones in the pituitary gland (also named hypophysis). However, some of the chromophobe cells may also be undifferentiated, nonsecretory cells [4, 15]. Glucocorticoids have several functions, including protein catabolism and stimulation of the hepatic gluconeogenesis from amino acids [10, 37]. Certain breeds, including Australian Terrier, Keeshounds, Alaskan Malamutes, Finnish Spitzes, Standard and Miniature Schnauzers, Miniature Poodles, and English Springer Spaniels, seem to be at increased risk to develop diabetes. The backup of the digestive juices elevates the pressure in the minor duct that causes abdominal pain and pancreatitis. Bar = 25 µm.Insulin and glucagon are pancreatic hormones that play pivotal roles in regulating glucose homeostasis and metabolism, which have opposite effects on glycaemia as well as on the metabolism of nutrients [24, 26, 28].
Others, such as Boxers, German Shepherd dogs, Cocker Spaniels and Collies seemed to be at decreased risk [40, 52].
So, the actions of the two hormones combined contribute to the control of blood glucose.Glucagon secretion by ?-cells is highly regulated by multiple factors, being the most important the glucose and insulin levels. Exercise intolerance or decreased activity, ketonic breath, recurrent infections (urinary tract, conjunctivitis), cataracts and hepatomegaly can also be present [45]. Other tests that can help diagnose pancreas divisum without the risk of causing pancreatitis include endoscopic ultrasound (EUS), and MRI (magnetic resonance imaging). Bar = 25 µm.The pars tuberalis is composed by cuboidal weakly basophilic cells, arranged in cords, nests or follicles and its function is not yet well established [4].
Low glucose levels activate specific channels in the brain and in pancreatic ?-cells to generate action potentials of sodium and calcium currents, leading to glucagon secretion. Bar = 25 µm.The zona reticularis is also composed of polyhedral cells, whose arrangement consists in freely anastomosing cords (Figure 19) [32]. Also, somatostatin inhibits glucagon secretion by inhibiting adenylate cyclase and cAMP production [26]. These cells contain less lipid content but have densely granular cytoplasm for which they are called “compact cells” [34].
Spontaneous hyperadrenocorticism is associated with inappropriate secretion of ACTH by the pituitary (pituitary-dependent hyperadrenocorticism) or with a primary adrenal disorder (adrenal-dependent hyperadrenocorticism) [40, 53, 54]. Insulin has several physiologic actions that include stimulation of cellular glucose and potassium uptake [29]. Over 80% of dogs with spontaneous hyperdarenocorticism suffer from pituitary dependent hypercorticism, resulting in an over secretion of ACTH [53, 54]. Some doctors will attempt sphincterotomy (cutting of the minor papilla during ERCP to enlarge its opening).
Glucose-induced insulin secretion from pancreatic ?-cells depends on mitochondrial activation, amongst other factors, like ATP, glutamate and others [30]. These hormones are produced in the hypothalamus, transported through the hypothalamic-pituitary axis and stored in the neurohypophysis, until a stimulus induces their release [4, 16] (Table 2).ADH has several effects on the body, particularly in terms of water saving and in increasing blood pressure.
Incretin hormones, glucose-dependent insulinotropic polypeptide (GIP) and glucagon like peptide-1 (GLP-1), secreted by cells of the gastrointestinal tract in response to meal ingestion, exert important glucorregulatory effects, including the glucose-dependent potentiation of insulin secretion by pancreatic ?-cells [31].Table 5 summarizes hormones of the pancreas and their functions.
Oxytocin stimulates contraction of the myoepithelial cells of the mammary gland, causing the ejection of milk. Adrenal-dependent hyperadrenocorticism represent the remaining 20% of spontaneous hyperadrenocorticism and is generally associated with unilateral (or occasionally bilateral) adrenal tumours. Therefore, patients with symptomatic pancreas divisum are best evaluated and treated by doctors in centers experienced in treating pancreatic diseases.
It also binds to the smooth muscle cells of the uterus, promoting uterine contractions during parturition [10] (Table 2).
Dogs showing pituitary-dependent hyperadrenocorticism exhibit a mean age of 7-9 years old, and those with adrenal-dependent hyperadrenocorticism, a mean age of 11-12 years old.
There are some breeds that are most frequently associated with hyperadrenocorticism, but any breed can develop it. The medulla consists of large columnar or polyhedral secretory cells, randomly distributed, with rich blood supply (Figure 20). The cells have large, vesicular nucleus, basophilic cytoplasm with fine positive chromaffin granules, due to the presence of catecholamines such as epinephrine and norepinephrine, which after exposure to oxidizing agents, such as chromate, yield a brown reaction by the formation of colored polymers [4, 31, 33]. This can arise from a destruction of more than 90% of both adrenal cortices, causing inability to produce corticosteroids (Primary hypoadrenocorticism or Addison’s disease), or from a deficiency in ACTH production by the pituitary (Secondary hypoadrenocorticism) [56, 57].
Besides different pathophysiology, primary and secondary hypoadrenocorticism also exhibit different clinical signs.
In adults, there are 3 types of adrenal medullary cells (1): epinephrine cells (66–75%), norepinephrine cells (25–33%) and small granule-containing cells (SGC, 1-4%). It is often associated with idiopathic adrenocortical atrophy, or with some therapeutic or surgical procedures, like mitotane treatment for hyperadrenocorticism or bilateral adrenalectomy.
The adrenal SGC cells of dogs vary from cells with a few granules and a high nucleo-cytoplasmic ratio to cells filled with many granules and a large mass of cytoplasm. This condition generally affects young and middle age dogs with a median age of 4 years [57].
Most of the chromaffin granules of these cells are small, ranging from 100 to 200 nm in diameter [39]. Great Danes, Portuguese Water Dogs, Rottweilers, Standard Poodles, West Highland White Terrier and Soft-coated Wheaten Terriers are the breeds in greater risk to develop hypoadrenocorticism [56].
The medullary cells produce other peptides in addition to epinephrine and norepinephrine, such as met-enkephalin, substance P, neurotensin, neuropeptide Y, and chromogranin A. The adrenal medulla contains also presynaptic sympathetic ganglion cells, randomly scattered [4, 32]. The first step in the synthesis of epinephrine is the enzymatic conversion of tyrosine to dihydroxyphenylalanine (DOPA) by tyrosine hydroxylase.
The animal can be in a hypovolaemic shock, and generally is found in a state of collapse, or collapses when stressed. Weak pulse, bradycardia, abdominal pain, vomiting, diarrhoea, dehydration and hypothermia can also be present. Norepinephrine then leaves the granule to be converted into epinephrine in the cytosol by phenylethanolamine-N-methyltransferase (PNMT), and epinephrine re-enters the granule for storage in the cell.
Animals with chronic hypoadrecocorticism may present anorexia, vomiting, lethargy, depression and weakness [57].8. The activity of PNMT is induced by the high local concentration of glucocorticoids in sinusoidal blood from the adrenal cortex [10, 32].
ConclusionsMalfunction of endocrine glands leads to severe multivariate syndromes, requiring a specialized medical approach and appropriate nursing care.
Acute stress, hypoglycemia or other similar situations, result both in catecholamine secretion and in transsynaptic induction of catecholamine biosynthetic enzymes, including tyrosine hydroxylase.
For this reason, professionals engaged in clinical veterinary practice need to know animal organic structures and how they function as a whole, including the role of the endocrine system, its glands and their hormones. Other environmental influences, including growth factors, extracellular matrix, and a variety of hormonal signals that generate cyclic AMP, also may regulate the function of chromaffin cells [32]. This systematic approach to endocrine disorders promotes not only a trained professional but a professional with technical and scientific knowledge, enabled with the exact notion of the involved etiopathogenic mechanisms and capable of making a diagnostic and therapeutical difference. In table 6, we can see a summary of some of the hormones produced by adrenal glands and their functions. This concerted attitude would certainly contribute to improve veterinary medical care to a level of excellence in the care of animal patients.NaN.
AcknowledgementsThis work was supported by Portuguese Foundation for Science and Technology and Center for Studies in Education, Technologies and Health.




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