Chapter 22
Avian Endocrine System
Colin G. Scanes
Department of Biological Sciences, University of Wisconsin, Milwaukee, WI, USA
22.1 INTRODUCTION
Hormones play a critical role in controlling many aspects
of the physiology of birds, including reproduction, metabo-
lism, and growth. Hormones are produced from ductless or
endocrine glands. They are then released into the blood-
stream. The hormones then act on specific tissues, cells, or
organs (referred to as the target organ). Each hormone acts
by interacting with receptors on the surface of the cells (pro-
teins and polypeptides) or within the cytoplasm and nucleus
(steroids and thyroid hormones). Birds have a complement
of endocrine organs similar to those of mammals:
l Anterior pituitary gland (discussed elsewhere in this vol-
ume in Chapter on the pituitary gland)
l Hypothalamus posterior pituitary gland complex (also
discussed in Chapter on the pituitary gland)
l Gonads (discussed in Chapters on male and female repro-
duction)
l Pancreatic islets (discussed in Chapter on the endocrine
pancreas)
l Adrenal glands (discussed in Chapter on the adrenal
glands)
l Thyroid glands (discussed in Chapter on the thyroid
glands)
l Parathyroid glands (discussed in Chapter on the role of
hormones in the regulation of bone turnover and eggshell
calcification)
l Pineal body or gland (discussed in Chapter on the pineal
gland)
l Ultimobranchial bodies or glands (C cells producing cal-
citonin). The ultimobranchial glands are found during
development in mammals, with the C cells then becoming
distributed in the thyroid and parathyroid glands. (This is
also discussed in Chapter on the role of hormones in the
regulation of bone turnover and eggshell calcification.)
l Endocrine cells of the gut (discussed in Chapter on the
gastrointestinal anatomy and physiology).
The traditional view of a distinct endocrine system, with
hormones only produced by specific endocrine glands that
Sturkies Avian Physiology.
Copyright 2015 Elsevier Inc. All rights reserved.
do little else, is overly restrictive. There is production of
biologically active proteins and peptides by many, if not all,
tissues. Organs that were thought of as simply target organs
for hormones also produce hormones such as the following:
l Adipose tissue producing adiponectin and leptin (dis-
cussed in Chapter on adipose tissue)
l Heart cells producing natriuretic peptides
l Liver producing insulin-like growth factor-1 (IGF1)
l Kidney producing renin (and hence angiotensin-I, then
angiotensin-II), 1,25-dihydroxy vitamin D3, and eryth-
ropoietin (discussed in Chapters on blood and osmoreg-
ulation)
22.2 AVIAN PHYLOGENY
Caution should be expressed about extrapolating studies for
poultry species such as chickens to other birds due to the
multiple generations of selection, with chickens domesti-
cated about 10,000 years ago (Sawai etal., 2010). There are
differences between the present poultry and the ancestral
stock. For instance, the thyroid-stimulating hormone (TSH)
receptor gene exhibits the same mutation throughout mul-
tiple lines of domesticated chickens but not in jungle fowl
(Rubin etal., 2010). Moreover, most of the coding sequence
of SH3RF2 is deleted (Rubin etal., 2010).
Even without domestication, it is argued that neither
galliform (e.g., chickens, turkeys, and quail) nor anseri-
form birds (ducks and geese) are necessarily good models
for other avian species. Within the class Aves (birds) and
Neoornithes (modern birds, including all extant species),
there are two branches:
1. Palaeognathae (including emus, ostriches, and rhea)
2. Neognathae, consisting of two distinct groups:
a. Galloanserae, consisting of the orders Anseriformes
and Galliformes
b. Neoaves (most modern species of birds) (see the Tree
of Life project, http://tolweb.org; and see Hackett
etal., 2008; Jetz etal., 2012; McCormack etal., 2013)
489
490
The two branches, Palaeognathae and Neognathae,
diverged about 140 million years ago (Sanders and
Lee, 2007), while the Galloanserae and Neoaves
separated about 95 million years ago (Ericson etal.,
2006). Within the Neoaves, there are three clades:
Water-birds
Land-birds
A third clade containing bustards (family:
Otididae), cuckoos (family: Cuculidae), flamingos
(family: Phoenicopteridae), hummingbirds (fam-
ily: Trochilidae), trumpeters (family: Psophiidae),
and turacos (family: Musophagidae), exhibiting
marked apparent diversity
Pigeons and doves (family: Columbidae) form a dis-
tinct side group within the Neoaves (Hackett etal., 2008;
McCormack etal., 2013).
22.3 PEPTIDES AND OTHER CHEMICAL
MESSENGERS CONTROLLING PHYSIOLOGY
The same chemical entity may act in one or more of the fol-
lowing manners:
l As a hormone traveling from endocrine cells to target
cells via the bloodstream
l As a paracrine factor released from one cell type to affect
another cell type passing through the extracellular fluid
l As an autocrine factor released from one cell type to
affect another cell of the same type passing through the
extracellular fluid
l As an intracrine factor acting with a cell type to affect
another cell
l As a neurotransmitter or neuromodulator released from a
neuron to act with another neuron
l As a neuropeptide released from a neuron to act with
other cell types in close proximity
l As a neuroendocrine factor released from neurosecretory
terminals of a neuron into the blood and then acting with
other cell types
Many of these chemical messengers are neuropeptides.
Table 22.1 summarizes a list of neuropeptides and protein
polypeptide hormones found in avian species. This is
based on the genomics of both the chicken and zebra finch
(Taeniopygia guttata) (Delfino etal. 2010; Xie etal., 2010).
Table 22.2 lists neuropeptides found in other vertebrate
groups but not in birds.
Throughout vertebrate evolution, there are multiple
examples of gene duplication leading to new chemical sig-
nal peptides and proteins or to new receptors. There are
also cases of gene loss. For instance, prolactin-like protein
(PLP) is a gene found in both fish and birds but not mam-
mals (Wang etal., 2010). Presumably, the gene was retained
in the amphibian and reptilian ancestors of birds but lost in
the ancestors of mammals.
PART | V Endocrine Theme
Our ability to understand the physiology of birds is
being facilitated with the tremendous progress in genom-
ics (discussed in detail in Chapter on genomics). It should
be noted that there can also be polymorphisms at the gene
level, splicing differences of RNA, and posttranslational
differences, including proteolytic cleavage (e.g., with the
translated products of the POMC and glucagon genes) and
glycosylation (e.g., with luteinizing hormone (LH), follicle-
stimulating hormone (FSH), and TSH).
22.4 CHEMICAL MESSENGERS FOUND IN
BIRDS BUT NOT MAMMALS
There are avian neuropeptides and their respective genes
that are not present in mammals. These include corticotro-
pin-releasing factor (CRF) amide, c-type natriuretic peptide
1 precursor, renal natriuretic peptide (Delfino etal., 2010),
and PLP (Wang etal., 2010).
22.5 HORMONES PRODUCED BY
NONTRADITIONAL ENDOCRINE ORGANS
22.5.1 Adiponectin
A nice example of a protein produced by organs not tradi-
tionally thought of as endocrine organs is the putative hor-
mone adiponectin. This is expressed in multiple tissues of
the chicken, including the adipose tissue (highest), liver (next
highest), anterior pituitary, brain, kidney, and skeletal mus-
cle (Maddineni etal., 2005). In addition, these tissues also
express the adiponectin receptors, AdipoR1 and AdipoR2,
with the highest expression of AdipoR1 being in skeletal
muscle, adipose tissue, and brain and with some expression in
anterior pituitary gland, kidney, liver, ovary, and spleen, and
with the highest expression of AdipoR2 being in adipose tis-
sue followed by skeletal muscle and some expression also in
anterior pituitary gland, brain, kidney, liver, ovary, and spleen
(Ramachandran etal., 2007). It can readily be envisioned that
adiponectin may act as a hormone and as a paracrine factor.
22.6 UNIQUE ASPECTS OF BIRDS
22.6.1 Song
Passerine birds have a unique ability for song. This involves
complex controls from the brain. There has been rapid
molecular evolution of the song-related genes in passerine
birds (Warren etal., 2010).
22.6.2 Salt Glands
Salt glands allow some birds to prosper in marine environ-
ments by secreting a hypertonic solution, predominantly
sodium chloride (discussed in detail in Chapter on osmoreg-
ulation). The salt glands respond to hormones. Angiotensin
Chapter | 22 Avian Endocrine System 491

TABLE 22.1 Neuropeptides and PeptideProtein Hormone in Birds

Pre-proneuropeptide or Official Gene Present in Chicken Present in Zebra Finch Other, Avian-Specific
Neuropeptide Symbol Genome?1 Genome?2 Notes
Adenylate cyclase activating Adcyap1 Expressed (e.g., in the
polypeptide 1 (pituitary) brain and gastrointesti-
nal (GI) tract)
Adrenomedullin AMD Expressed in lungs,
adrenal glands, skeletal
muscle, and GI tract
(Zudaire etal., 2005)
Expression increased
in lungs of hypotensive
chicken (Gomez etal.,
2007)
Arginine vasopressin AVP Neuropeptide in birds
arginine vasotocin
(AVT)
C-RF amide peptide LOC420716/
CRF
C-type natriuretic peptide 1 CNP 1 Expressed in the
chickendiscussed in
narrative
C-type natriuretic peptide 3 CNP 2
Calcitonin-related Polypeptide alpha CALCA
[also called calcitonin gene-related
peptide (CGRP)] Gene encodes
calcitonin also
Cocaine- and amphetamine-regulated CART X CNS effect in chickens
transcript protein (Tachibana etal., 2003)
Corticotropin-releasing hormone CRH Expressed in
(also referred to as corticoliberin or hypothalamus
corticotropin-releasing factor (CRF))
Cholecystokinin CCK
Chromogranin A (parathyroid CHGA
secretory protein 1)
Chromogranin B CHGB
Chromosome 12 open reading frame C12orf39
39 (spexin)
Chromosome 2 open reading frame 40 C2orf40
Endothelin 1 EDN1 Expression increased
in lungs of hypotensive
chicken (Gomez etal.,
2007)
Endothelin 2 EDN2
Endothelin 3 EDN3
Follicle stimulating hormone beta FSHB Expressed in adenohy-
subunit pophyseal cells
Galanin pre-propeptide GAL Expressed in CNS and
GI tract

Continued

492 PART | V Endocrine Theme

TABLE 22.1 Neuropeptides and PeptideProtein Hormone in Birdscontd

Pre-proneuropeptide or Official Gene Present in Chicken Present in Zebra Finch Other, Avian-Specific
Neuropeptide Symbol Genome?1 Genome?2 Notes
Gastric-inhibitory polypeptide (GIP) GIP Expressed in CNS and
GI tract
Gastrin-releasing peptide (GRP) GRP Expressed in GI tract
and hypothalamus
Ghrelin or obestatin pre-propeptide GHRL Expressed in GI tract
and hypothalamus
Glucagon pro-hormone also GCG Expressed in pancreas,
containing glucagon-like peptide 1 GI tract, and
(GLP1) and GLP2 hypothalamus
Glycoprotein hormone, polypeptide CGA Expressed in
adenohypophyseal cells
Gonadotropin-releasing hormone GnRH Expressed in hypothala-
(GnRHI) (formerly luteinizing mus and second gene
hormonereleasing hormone I encoding GnRHII in
(LHRHI)) birds
Growth hormone GH Expressed in adeno-
hypophyseal cells and
CNS
Duplicate GH gene on
chromosome 1 on
passerine birds
Expression of duplicate
gene not well
established
Growth hormonereleasing hormone GHRH Expressed in
(GHRH), also called somatoliberin hypothalamus
Hypocretin (orexin) neuropeptide HCRT Expressed in CNS
precursor
Insulin INS Expressed in pancreas
Insulin-like 5 INSL5 X
Insulin like growth factor 1 IGF1 Expressed in liver and
cartilage
Insulin like growth factor 2 IGF2
Islet amyloid polypeptide IAPP Functionally similar
to calcitonin-related
polypeptide alpha;
expressed in CNS, GI
tract, and pancreas
(Fan etal., 1994)
Kisspeptin KISS1 X X Effects of kisspeptide
10 are reported in the
chicken ovary
(Xiao etal., 2011).
Leptin or leptin-like OB ? See narrative in text for
details
Luteinizing hormone beta subunit LHB Expressed in adenohy-
pophyseal cells

Chapter | 22 Avian Endocrine System 493

TABLE 22.1 Neuropeptides and PeptideProtein Hormone in Birdscontd

Pre-proneuropeptide or Official Gene Present in Chicken Present in Zebra Finch Other, Avian-Specific
Neuropeptide Symbol Genome?1 Genome?2 Notes
Mesotocinneurophysin 1 MST Equivalent to oxytocin
in mammals
Expressed in CNS
Motilin MLN Expressed in CNS and
GI tract
Natriuretic peptide precursor A NPPA
Neuromedin B NMB
Neuromedin U NMU
Neuropeptide S NPS
Neuropeptide VF (gonadotropin-inhibi- NPVF
tory hormone (GnIH))
Neuropeptide W NPW X
Neuropeptide Y NPY
Neurotensin NTS Expressed in CNS and
GI tract
Osteocrin OSTN
Pancreatic polypeptide PPY (or PP) Expressed in pancreas
and GI tract
Parathyroid hormone PTH Expressed in
parathyroid gland
Parathyroid hormonelike PTHLH Expressed in early
hormone (also called embryonic develop-
PTH-related peptide (PTHrP)) ment skeletal tissues
For expression, also see
Chapter on the pituitary
gland
Pituitary adenylate cyclaseactivating See adenylate cyclaseactivating polypeptide
polypeptide 1
Platelet-derived growth factor D PDGFD
Platelet-derived growth factor alpha PDGFA
polypeptide
Platelet-derived growth factor beta PDGFB
polypeptide
Pre-pronociceptin PNOC
Pro-melanin-concentrating PMCH
hormone (MCH)
Prodynorphin PDYN
Proenkephalin PENK CNS
Prokineticin 2 PROK
Prolactin PRL Adenohypophyseal
cells
Prolactin B (also called prolactin-like PRLB Reported in birds and
protein (PLP)) fish but not mammals

Continued

494 PART | V Endocrine Theme

TABLE 22.1 Neuropeptides and PeptideProtein Hormone in Birdscontd

Pre-proneuropeptide or Official Gene Present in Chicken Present in Zebra Finch Other, Avian-Specific
Neuropeptide Symbol Genome?1 Genome?2 Notes
Prolactin-releasing hormone PRH CNS
Pro-opiomelanocortin POMC CNS and adenohy-
pophyseal cells
Pyroglutamylated RF amide
Relaxin 3
Secretin CNS and GI tract
Secretogranin II (chromogranin C)
Secretogranin V (7B2 protein)
Somatostatin SST1 CNS, GI tract, and
pancreas
Somatostatin 2 SST2
Tachykinin, precursor 1 TAC1
Thyroid-stimulating hormone TSHB Adenohypophyseal
(or thyrotropin) beta subunit cells
Thyrotropin-releasing hormone TRH CNS
(also called thyroliberin)
Urocortin UCN
Urocortin 3 UCN3
Urotensin 2 UTS2
Urotensin 2 domain-containing UTS2D
Vascular endothelial growth factor C VEGFC
Vasoactive intestinal peptide VIP CNS and GI tract
1Based on Delfino etal., 2010.
2Based on Xie etal., 2010.

II directly decreases fluid flow from the duck salt gland
(Butler, 2007). This effect is independent both of the adrenal
gland and of catecholamines as angiotensin II is effective in
adrenalectomized and chemically sympathectomized birds
(Butler, 2007). Natriuretic peptides influence the function-
ing of the salt glands (Schtz and Gerstberger, 1990).
22.6.3 Unique Aspects of Metabolism
There are marked differences in circulating concentrations of
glucose between wild birds and mammals, with avian con-
centrations double those in mammals and with physically
smaller species of birds or mammals having higher circulat-
ing concentrations of glucose than larger ones (Braun and
Sweazea, 2008) (discussed in detail in Chapters on carbo-
hydrate metabolism and the pancreas). Moreover, birds do
exhibit diabetes, with circulating concentrations of glucose
held at a consistent level, except under one paradigm. Chick-
ens that received passive immunization with antisera to insulin
exhibit diabetic-like shifts in metabolism, with circulating
concentrations of glucose of 747mg/dL reported 5h after
antisera administration (Dupont etal., 2008; Simon etal.,
2012). Interestingly, identical extremely high concentrations
of glucose are observed following feeding in hummingbirds
without adverse effects (Beuchat and Chong, 1998).
Birds also have high metabolic demands for both short-
term flight and extended migratory flight (discussed in
detail in Chapter on migration).
22.6.4 Reproduction
Birds exhibit marked differences from mammals in terms of
reproduction (discussed elsewhere in this volume in Chapters
on male and female reproduction). These differences include
the internal testes, the production of large yolky eggs, the
absence of pregnancy, the existence of only one ovary and
oviduct (the Mllerian duct), and unique sexual, brooding,
and parental behavior (discussed in Chapter on brooding).
Chapter | 22 Avian Endocrine System 495

from the cDNA (Liu etal., 2007). The leptin receptor is

TABLE 22.2 Neuropeptides Presently Not present in multiple tissues, including the brain and liver
Demonstrated to Exist in Birds (Based on Delfino etal., in the chicken, based on expression (Liu etal., 2007)
2010) and the observed immunoreactive 180kDa protein in
Apelin (APEL) Proenkephalin-B (PDYN) multiple tissues (Ohkubo etal., 2007). The leptin recep-
Cortistatin (CORT)1 Peptide YY (PYY)
tor is expressed in the anterior pituitary gland and basal
hypothalamus of the chicken (Liu and Sharp, 2007). The
Galanin-like peptide (GALP) Putative peptide YY-2 leptin receptor has been characterized also in the turkey
(PYY2)
(Richards and Poch, 2003) and goose (Wang etal., 2011).
Hepcidin (HEPC) Pro-relaxin 1 (REL1) The leptin receptor is expressed in the brain, lung, and
Insulin-like 3 (INSL3) Pro-relaxin 2 (REL2) spleen, with some expression also in the adipose tissue,
duodenum, liver, pancreas, and skeletal muscle (Richards
Insulin-like 4 (INSL4) Regulated endocrine-
specific protein 18 (RES18)
and Poch, 2003; Wang etal., 2011). Adipose expression
of the leptin receptor is increased by oleic acid invitro
Intermedin (ADM2) Spexin (SPXN) (Wang etal., 2011). In ovo administration of mammalian
Metastasis-suppressor KiSS-1 Tachykinin, precursor 2 leptin into the albumin of chick embryos was reported
(KISS1)2 (TKN2) to result in increased posthatching growth and circulat-
Natriuretic factor B (ANFB) Tachykinin, precursor 3 ing concentrations of triiodothyrone (T3), with greater
(TKN3) hepatic expression of growth hormone receptor and
Neuromedin-S (NMS) Tachykinin, precursor 4
IGF1 mRNA (Li etal., 2011). Invitro mammalian leptin
(TKN4) elevates expression and activities of deiodinase in chick
embryo hepatocytes (Li etal., 2011). Administration of
Neuropeptide B (NPB) Torsin family 2, member
A (TOR2X)
mammalian leptin decreases food intake and fat mass in
wintering white-throated sparrows (Cerasale etal., 2011).
Neuropeptide FF (NPFF) The gene for the leptin receptor has been identified in the
Neuropeptide S (NPS) zebra finch (T. guttata) (leptin receptor (LEPR) Gene ID:
Proprotein convertase subtilisin/
100229897).
kexin type 1 inhibitor (PCSK1N)
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