Biology 102: Comparative Anatomy of Vertebrates

Synthesis Paper
Gladys Ericka Galang
Pancreas: General Form and Function in Avian Species

The avian pancreas is located on the right side of the abdominal cavity (Mobini, 2013) in the
duodenal loop (Rideau, 1988) in all birds and is the largest gland connected with the
alimentary tract. It consists of the endocrine portion or pancreatic island and the exocrine
portion (Mobini, 2013) where more endocrine tissues than exocrine tissues. Despite this,
99% percent of the mass of the avian pancreas has exocrine functions. (Doneley, 2001)
Three lobes comprise the gland: the dorsal and ventral lobes which represent a large part of
the gland, and the cranial splenic lobe that is smaller than the two. (Rideau, 1988) Within the
islands are islets that hold cells and acini that secrete hormones and enzymes that are
responsible for digestion and homeostasis, among other functions of the pancreas. (Mobini,
2013)
Although the pancreas also functions in digestion, this synthesis paper will focus on the
endocrine functions of the pancreas, and therefore, the pancreatic island.
The endocrine pancreas in general come in the form of clusters of cells called the Islets of
Langerhans. (Bowen, 2002) The islets are found more frequently in the tail part of the whole
pancreas than in the head and the body. (Carlos, 2007) Avian pancreas contains two islet
types: alpha, and beta islets that contain cells (Mobini, 2013) that release hormones for,
mostly, glucose regulation. (Sitbon et al, 1980) and carbohydrate metabolism. The pancreas
of some species, like that of the mature pigeons, Columba livia, contain only these two islets
while other species, such as the ducks, contain a mix of the two cells in one (Mobini, 2013),
and even some others like the Australian wedge-tailed eagles, Aquila audax, only have the
mixed type (Edwin et al, 1993).
The splenic lobe contains the highest percentage of islet tissues (Guha et al, 1978) with high
concentration of the different islet cells (Edwin et al, 1993) and contains the largest islets
while the ventral lobe contains islets of intermediate and the dorsal lobe contains the
smallest islets, as found consistently in 13 different species of Indian birds (Guha et al,
1978).

Alpha islets contain alpha and beta cells, and beta cells contain beta and delta cells (Mobini,
2013) Alpha-cells which are morphologically denser than beta cells in the pancreatic tissue
as found by immunohistochemistry (Hostetter, 2010) release glucagon while beta cells
release insulin (Hazelwood, 1973). Insulin and glucagon work antagonistically in maintaining
blood sugar balance, in all vertebrates (http://biomed.brown.edu/). Delta cells, also found in
different parts of the digestive system, release somatostratin which generally functions to
inhibit the secretion of other hormones of the body. (Bowen, 2002) Alpha cells are found
most abundantly in the Indian bird species, with relative abundance of delta cells, (Guha et
al, 1978) while it was found that beta cells are most numerous in the pancreas of the
Brazilian sparrow. (Carlos, 2007) Alpha islets are smaller than beta cells with 2-3 cells, while
the beta cells are clearer and larger in the zebra finch (Saadatfara, et al, 2010). Compared to
mammals, the endocrine pancreas of birds contain similar cell populations similar with
minimal differences in the distributions of these cell types (Hazelwood, 1983)
In Hazelwoods (1993) article, he introduces two kinds of alpha cells, A1 and A2, which are
found in alpha islets. He claims beta islets contain beta and A2 cells and that A2 cell release
glucagon and beta cells release insulin, while the functions, specifically the hormones
released by A1 cells are uncertain.
A number of changes of the islets of Langerhans can be found during sexual activity of birds
of both sexes and during the egg-laying season of female birds particularly the pigeons
where the pancreatic islets generally increase in size and number. (Schulz, 1940)
There are subtle differences found in the pancreas of the avian species. Carlos (2007)
observed that beta cells, which he called the insulin-immunoreactive cells, are most
numerous and occupies the central portion of the islets in the pancreas of the Brazilian
sparrow. The alpha cells and delta cells which he referred to as glucagon-immunoreactive
cells and somastotratin-immunoreactive cells are found at the periphery and interior portion
of the islets.
A comparison between the pancreas of the zebra finch, Poephila bichenovi and pigeon
Streptopelia decaocto was made by Saadatfara, et al, (2010). The two birds are mostly
similar with the major difference lying on the number of cells found in the lobes.They found
that the ventral lobe contained a greater number of islets than the beta islets, and a
dominance of the alpha cells. The splenic lobe contains more beta cells but without the
absence of the alpha cells. The pigeons, however, as agreed by the study of Mobini (2013)
with a different species, have pancreas with beta cells more numerous than the alpha cells.
A great number of islets were distributed in a great area of the splenic lobe. The ventral lobe

contains irregularly-shaped alpha islets while the dorsal lobe contains regularly-shaped beta
islets.
In another study of pigeons, the pancreas is described to be a serous tubuloacinar gland
covered by a thin collagenous capsule that was found also in geese but was found thick in
turkeys. There were no major differences found based on the sex of the avian species.
Septa from connective tissue of the capsule divide the pigeon pancreas into indistinct lobes
and lobules, unlike the previously mentioned species with clear ventral, dorsal, and splenic
lobes. Instead, light and dark islets were found in pigeons. (Schulz, 1940) Parasympathetic
ganglia were found in the interlobular connective tissues as were found in different studies of
ducks, turkeys, and chickens. (Mobini, 2013)
Endocrine pancreas generally function for the synthesis and secretion of hormones in
maintaining balance (Bowen, 2002), mostly in the blood glucose level or plasma glucose
PGlu. Some of the hormones were already previously discussed: insulin which increases the
affinity of organs and cells to glucose, glucagon which stimulates the increase of glucose in
the bloodstream, and somastotratin which inhibits the secretion of the two. Other hormones
include the avian pancreatic polypeptide, and YY polypeptide (Carlos, 2007). Avian
pancreatic secretion is found to have a higher rate than in mammals, controlled by both
neural and hormonal mechanisms (Doneley, 2001)
Plasma glucose concentration in birds is higher than in mammals with similar boy mass.
(Downs et al, 2009) The broiler chicken, Gallus domesticus, has a PGlu level that is twice as
high as in humans and four times as high as in mammals with equal body mass even with
the same insulin concentration which is the consequence of development of insulin
resistance in post-hatch life, and perturbations in the insulin signalling cascade (Franssens
et al, 2014). Pancreatic insulin concentration lies between 20 and 80 ng/mg tissue in
chickens and pigeons and between 1.5 and 4ng/mg tissue in the pancreas of the duck.
(Rideau, 1988)
Studies have also found that the avian pancreas contains small amounts of insulin, and
releases the hormone with inefficient speed in response to high glucose load. (Hazelwood,
1973) Pancreas of 4-8 week old broiler chickens had been found to be relatively insensitive
to initiatory fuel nutrients such as D-glyceraldehyde, D-mannose, L-leucine, and alphaketoisocapoic acid, at concentrations which induce a maximal insulin release in mammalian
pancreas. High glucose concentrations are required to get a two-fold increase in insulin
secretion from pancreatic islets of 5-6-day old Rhode Island Red chicks, compared to the

normal low glucose concentration is already sufficient for pancreatic secretion in mice.
(Franssens et al, 2014)
Fasting, hormones, and/or vagal stimulation do not influence insulin release. (Hazelwood,
1973) Some studies have concluded that the PGlu concentration is insensitive to the
regulation of insulin in the blood. (Braun et al, 2008) since it was found that insulin was not
significantly correlated with variations in PGlu, (Calhoun, n.d.) which may be cause by the
absence of the glucose transporter GLUT4 that is found in mammals. (Franssens et al,
2014) Interestingly enough, high levels of PGlu don't lead to significant oxidative stress to
birds as it is known that they are longer-lived compared to mammals. (Braun et al, 2008)
The avian hormones that are in charge of glucose regulation are glucagon, insulin,
corticosteroid, thyroxine, pancreatic polypeptides (APP) with corticosterone being the main
glucocorticoid. The function in the removal of glucose for normalization after increase
because of post-digestion and absorption concentration and glucose synthesis by
gluconeogenesis and glycogenolysis. (Downs et al, 2009)
More than 70% of glucose in blood stream is made available by the liver where transition to
gluconeogenesis is rapid and continuous by using amino acids as substrata as found in
some carnivorous species (Downs et al, 2009) which may be caused by high levels of APP
that also induces hypoglycerolemia. (Hazelwood, 1973). Most birds store little glucose as
glycogen (Braun et al, 2008), so there is little glycogenolysis with higher proportions of
gluconeogenesis that lead to high glucose concentration in the blood. Primary substrates for
gluconeogenesis are glycerol and amino acids. (Downs et al, 2009)
Both avian pancreas and plasma contain 5-10 times more glucagon than observed in
mammals. Birds exhibit rapid hyperglycemia, hepatic glycogenolysis, hyperglycerolemia, and
hypertriglyceridemia upon treatment of mammalian glucagon (Hazelwood, 1973). Moreover,
immunohistochemical data from doves and quail show higher glucagon expression as
compared to insulin, which may contribute to their naturally high PGlu. (Calhoun, n.d.)
Glucagon appears to be a major pancreatic hormone involved in metabolic glucoregulation
in birds. (Hazelwood, 1983) Insulin may initially be the primary regulator of carbohydrate
metabolism during embryonic stage whereas glucagon may replace insulin after hatch
(Franssens et al, 2014). But huge considerations are made towards the molar ratio of insulin
to glucagon (I/G) as a dominant force in homeostasis rather than considering the two
hormones separately. (Hazelwood, 1983)

Another study had shown that PGlu concentration may be because of circadian variations.
Gender, feeding habits, feeding frequencies and captivity may also influence PGlu although
most data reveal consistency with little variations. Results show that fluctuations do occur
because of differences of temperature, with higher PGlu concentrations in lower temperatures
as was found in the Malachite sunbird Nectarinia famosa. They also found out that the
carnivorous birds had lesser basal PGlu concentration than the nectarivorous birds. (Downs
et al, 2009) PGlu regulation was also found to be age dependent, where embryonic chicken
have lower PGlu concentration than adults. (Franssens et al, 2014)
The high PGlu concentration is found to have developed because of high insulin production in
the embryonic stages of the chicken. Embryonic pancreatic insulin of white leghord chicken
increases concomitant to the development of the pancreas. Insulin displays a good response
to glucose level during the second and third week of embryonic development. The amount of
insulin elevates and peaks at the 17th embryonic day and plateaus during pipping and
hatching. Afterwards, sensitivity to insulin decreases in the growing chicken and finally the
adult broiler and layer chicken is regarded being insulin resistant because of differences in
the insulin signalling cascade in tissues and nervous system (Franssens et al, 2014).
Molar ratios between insulin and glucagon is indicative of a catabolic recovery of nutrients in
mammals, the ones found in the experiments suggest that birds normally are in a catabolic
mode. (Hazelwood, 1983) Arousal in birds require high energy costs, which may be the
reason of their high glucose concentration stored during torpor. Their carbohydrates are
consumed quickly during flight and foraging, and fats are substrata in fasted resting birds.
(Downs et al, 2009) Glucose is absorbed by the avian gastrointestinal tract by sodium
glucose co-transporters SGLT and upregulated. The avian nervous system utilizes glucose
as metabolic substrate. (Braun et al, 2008)
Diseases in the pancreas include pancreatitis and diabetes. Aetiology of pancreatitis found
in parrots is difficult to pinpoint. Pancreatitis generally develops in the activation of digestive
enzymes within the gland that results in pancreatic autodigestion. Pancreatic cell wall
damage releases the enzymes in the intracellular spaces and causes production of
unopposed free radicals that causes more enzymes to be released. Symptoms include
abdominal pain or gastro-intestinal dysfunction although these are not found in all birds with
pancreatitis, vomiting, diarrhea, ileus, anorexia, lethargy, weight loss, polyuria, polydypsia,
and abdominal distension. Factors that may contribute to pancreatitis are obesity, high fat
diet, zinc mycotoxin, selenium trauma, viral infection, Chlamydophila infection, bacterial
infection, egg yolk peritonitis, and neoplasia. But confirmation still is needed through
pancreatic biopsy. Known treatments are oral, subcutaneous, intravenous fluid therapy to

correct dehydration and improve perfusion. Surviving birds must be monitored for pancreatic
insufficiency and diabetes mellitus that follows most of mammalian pancreatitis patients.
(Doneley, 2001)

References
Bowen R. (2002) Functional Anatomy of the Endocrine Pancreas. Retrieved November 30,
2014, from http://arbl.cvmbs.colostate.edu/hbooks/pathphys/endocrine/pancreas/anato
Bowen, R. (2002). Somatostatin. Retrieved November 30, 2014, from
http://arbl.cvmbs.colostate.edu/hbooks/pathphys/endocrine/otherendo/somatostatin.html
my.html
Braun, E., & Sweazea, K. (2008). Glucose Regulation in Birds. Comparative Biochemistry
and Physiology Part B: Biochemistry and Molecular Biology, 151(1), 1-9.
Calhoun, M., McGraw, K., & Sweazea, K. (n.d.). Variations in pancreatic glucoregulatory
hormones in birds.
Carlos, R. (2007). Immunocytochemical study of the distribuition of endocrine cells in the
pancreas of the Brazilian sparrow species Zonotrichia Capensis Subtorquata. Brazilian
Journal

of

Biology,

67(4).

Retrieved

November

28,

2014,

from

http://dx.doi.org/10.1590/S1519-69842007000400021
Doneley, R. (2001). Acute Pancreatitis in Parrots. Australian Veterinary Journal, 79(9).
Retrieved November 26, 2014.
Downs, C., Wellmann, A., & Brown, M. (2009). Diel Vairiations In Plasma Glucose
Concentrations Of Malachite Sunbirds Nectarinia Famosa. Retrieved November 27, 2014.
Edwin, N., & Leigh, C. (1993). The endocrine pancreas in the Australian wedge-tailed eagle
(Aquila audax)--an immunocytochemical study. European Journal of Histology, 37(3), 219224.
Franssens, L., Buyse, J., Decuypere, E., & Everaert, N. (2014). Relationship Between
Glucose And Pancreatic Hormones During Embryonic And Posnatal Phase In Chickens.
Avian Biology Research, 7(1). Retrieved November 27, 2014.
Guha, B., & Ghosh, A. (1978). A cytomorphological study of the endocrine pancreas of some
indian birds. General and Comparative Endocrinology, 34(1), 38-44.
Hazelwood,

R.

(1973).

The

Avian

Endocrine

Pancreas. American

Society

of

Zoologists, 13(3), 699-709. Retrieved November 28, 2014.

Hazelwood, R. (1983). Pancreatic Hormones, Insulin/Glucagon Molar Ratios, and
Somatostatin as Determinants Of Avian Carbohydrate Metabolism. Journal of Experimental
Zoology, 232 (3), 647-52.

Hostetter, K., Sweazea, K., & Braun, E. (2010). The Avian Pancreatic Islets: A Comparison
across Species. Retrieved November 27, 2014.
Mobini, B. (2013). Histochemical and Histological Studies on the Pancreas of Mature Pigeon
Columba livia. European Journal of Experimental Biology, 3(2), 148-152. Retrieved
November 28, 2014.
Rideau, N. (1988). Chapter 25 Insulin secretion in birds. Leanness in Domestic Birds:
Genetic, Metabolic and Hormonal Aspects, 269-294.
Saadatfara, Z., Asadiana, M., & Alishahia, E. (2010). Structure of Pancreas in Zebra Finch
(Poephila bichenovi) and Pigeon (Streptopelia decaocto laughing nove). Journal of Applied
Animal Research, 37(2).
Schulz, H. (1940). The pancreas during the sexual cycle of the pigeon. Endokrinologie, 22,
319-330.
Insulin/Glucagon.

(n.d.).

Retrieved

November

30,

2014,

from

http://biomed.brown.edu/Courses/BI108/BI108_2002_Groups/pancstems/stemcell/insulin_gl
ucagon.htm