ArticlePDF Available

Morphological structure of the syrinx in the Bursa Roller Pigeon (Columba livia)

Authors:
Hüseyin Yıldız at Uludag University
Hüseyin Yıldız
Bestami Yilmaz at Harran University
Bestami Yilmaz
Ilker Arican at Uludag University
Ilker Arican
Download full-text PDF
Read full-text
Download citation

Abstract and Figures

The aim of this study was to investigate the topographical, anatomical and histological characteristics of the syrinx in 18 Bursa Roler Pigeons. The study showed that the syrinx in pigeons was of tracheobronchial type and was composed of tympanum, cartilagines tracheosyringeales, and cartilagines bronchosyringeales. Tympanum and cartilagines bronchosyringeales were formed from 5 cartilage rings, while cartilagines tracheosyringeales were formed from 4 C-shaped cartilage rings. The pessulus was made up of a double-folded mucous membrane extending dorsoventrally from median walls of the bronchus primarius into the cavum syringis. The membrana tympaniformis lateralis filled the distance between the third and fourth cartilagines tracheosyringeales rings. The membrana tympaniformis medialis covered the distance between the medial ends of the cartilagines broncho-syringeales rings. There were 2 syringeal muscles termed tracheolateral and sternotracheal. The lamina epithelialis of the mucosa was lined with non-cornified stratified squamous epithelium.
Figures - uploaded by Bestami Yilmaz
Author content
All figure content in this area was uploaded by Bestami Yilmaz
Content may be subject to copyright.
Content uploaded by Bestami Yilmaz
Author content
All content in this area was uploaded by Bestami Yilmaz on Jun 03, 2015
Content may be subject to copyright.
Bull Vet Inst Pulawy 49, 323-327, 2005
MORPHOLOGICAL STRUCTURE OF THE SYRINX IN THE
BURSA ROLLER PIGEON
(COLUMBA LIVIA)
HÜSEYIN YILDIZ
1
, BESTAMI YILMAZ
2
AND İLKER ARICAN
1
1
Department of Anatomy, Faculty of Veterinary Medicine, University of Uludag, 16059 Bursa, Turkey
2
University of Harran, 63300 Şanlıurfa, Turkey
e-mail: yildiz@uludag.edu.tr
Received for publication July 06, 2005.
Abstract
The aim of this study was to investigate the
topographical, anatomical and histological characteristics of
the syrinx in 18 Bursa Roler Pigeons. The study showed that
the syrinx in pigeons was of tracheobronchial type and was
composed of tympanum, cartilagines tracheosyringeales, and
cartilagines bronchosyringeales. Tympanum and cartilagines
bronchosyringeales were formed from 5 cartilage rings, while
cartilagines tracheosyringeales were formed from 4 C-shaped
cartilage rings. The pessulus was made up of a double-folded
mucous membrane extending dorsoventrally from median
walls of the bronchus primarius into the cavum syringis. The
membrana tympaniformis lateralis filled the distance between
the third and fourth cartilagines tracheosyringeales rings. The
membrana tympaniformis medialis covered the distance
between the medial ends of the cartilagines broncho-
syringeales rings. There were 2 syringeal muscles termed
tracheolateral and sternotracheal. The lamina epithelialis of the
mucosa was lined with non-cornified stratified squamous
epithelium.
Key words: pigeons, Columba livia, syrinx,
anatomy.
The Bursa Roller Pigeon lives in the South
Marmara Region in Turkey. Special attention has been
paid by breeders to protect the pureness of this race.
Important distinguishing feature of this race is their
fastidious vocal organ, called syrinx, allowing them to
produce their special song pattern (20, 13, 15). Its
morphological structure has been investigated in some
bird species. It is located in the terminal part of the
trachea and the first part of the two main bronchi (6, 10,
11, 16, 23, 25). The skeleton of the tracheobronchial
type of the syrinx is composed of 3 different parts, i.e.
tympanum, cartilagines tracheosyringeales (CTS) and
cartilagines bronchosyringeales (CBS) (2, 15, 16, 24).
Two membranes, membrana tympaniformis
lateralis (MTL) and membrana tympaniformis medialis
(MTM), are involved in the formation of sound in the
bird syrinx (13). The sound can be altered by a complex
of extrinsic and intrinsic muscles surrounding the syrinx
(16). The syrinx mucosa consists of either bistratified
squamous (1), columnar (3, 5, 17), or pseudostratified
prismatic epithelium (22).
In our knowledge, up to date no scientific study
has been carried out to characterize the structure of the
syrinx in the Bursa Roller Pigeon. In the present study,
we investigated the topography, anatomy, and histology
of the syrinx in this species. The results of this research
may useful in operations of the syrinx in this species.
Material and Methods
Eighteen (9 male and 9 female) Bursa Roller
Pigeons (Columba livia), which died due to various
causes, were obtained from the South Marmara Region
pigeon raisers. All the pigeons were 2-year-old and had
an average weight of 234.40 ± 11.12 g. Firstly, the body
cavity was opened and the syrinx was observed in the
terminal part of the trachea. After pointing out the
topographic features precisely, sections were made 6 cm
to the dorsal side of the trachea and 4 cm to the ventral
side of the syrinx. The morphological characteristics
were investigated by means of a magnifying glass and
measurements made with a digital compass (Mitutuyo
Corporation, Kawasaki, Japan). For histological
investigation, the tissue was fixed in 10% formalin for at
least 72 h. After routine processing and embedding in
paraffin, 6 µm tissue sections were cut, stained with
haematoxylin and eosin, and 10 µm sections were cut
and stained with Pincus acid orcein-Giemsa for the
demonstration of the elastic fibers surrounding the
cartilages (17). Sections were investigated under a light
microscope (Nikon Optiphot-2, Nikon Corporation
Imaging Company, Japan) and photographed (Nikon
FA-35DX, Nikon Corporation Imaging Company,
Japan).
Statistical analysis was performed by Minitab
statistical package (for Windows version 11). The
ANOVA was used to investigate comparison of the
means.
For the terminology, the Nomina Anatomica
Avium (14) was used.
324
Results
The pigeon syringes were observed to lie at the
dorsal side of the glandular stomach, ventrally of the
oesophagus and between the terminal part of the trachea
and bronchus primarius (Fig. 1). On the ventral surface
of the syrinx displayed itself the membrana
tracheosyringealis associated with the thin tympanum
and CTS in the midline. This membrane was not seen on
the dorsal surface of the syrinx. The skeleton of the
syrinx was composed of 3 different cartilage groups.
These were the tympanum, CTS and CBS (Fig. 2). The
results of the measurements of the laterolateral (LLD)
and dorsoventral (DVD) diameters of these parts are
presented in Table 1.
Fig. 1. Ventral view of the syrinx in the pigeon within the body cavity. Syrinx (s),
oesophagus (e), pessulus (p), membrana tympaniformis medialis (m).
Table 1
Mean laterolateral and dorsoventral diameters of the syringeal cartilages (mm)
First ring Third ring Fourth ring Fifth ring
LLD
5.10 ± 0.15
-
-
5.53 ± 0.12
*
Tympanum
DVD
3.64 ± 0.11
-
-
4.03 ± 0.09
*
LLD
5.67 ± 0.13
5.87 ± 0.15
5.22 ± 0.12
-
*
CTS
DVD
4.33 ± 0.14
5.03 ± 0.14
4.53 ± 0.10
-
**
LLD
2.36 ± 0.06
-
-
1.94 ± 0.06
**
CBS
DVD
4.12 ± 0.15
-
-
3.25 ± 0.12
**
* P< 0.05; ** P < 0.001; ± SD.
325
Fig. 2. Dorsal view of the pigeon syrinx. Tympanum (t), Fifth tympanum ring (t5), cartilago tracheosyringealis 3 (ts3),
cartilago bronchosyringealis (bs), cartilago bronchosyringealis 1 (bs1), membrana tympaniformis lateralis (l),
membrana tympaniformis medialis (m), m. sternotrachealis (st), m. tracheolateralis (tl).
Fig. 3. Histological section of the pigeon syrinx. Non-cornified stratified squamous epithelium (e),
mucous gland (g), lamina propria (lp), elastic fibres (el), hyaline cartilage (h), tunica muscularis (tm).
Pincus acid orcein-Giemsa stained, bar 50 µm.
326
The tympanum was formed from 5 oval-shaped
cartilage rings pressed dorsoventrally. Because of its
cylindrically formed cartilages, it was different from
CTS. The average length of the tympanum was 4.11 ±
0.09 mm and from cranial to caudal, both LLD and
DVD of these rings were increasing. In fact, the CTS
which are generally known as a part of the tympanum, in
our case appeared as a separate part. Ventral and dorsal
ends of CTS were not joined each other. The
composition of CTS was constituted from four cartilage
rings having an average length of 5.15 ± 0.13 mm. The
CTS were ovally shaped and its LLD were larger than
DVD. But, the diameter of the third cartilage was larger
than that of the fourth cartilage. The largest distance was
1.72 ± 0.06 mm in middle and 2.84 ± 0.09 mm in the
lateral part. The space between the third and fourth ring
was filled by a membrane known as MTL. The CBS
consisted of five C-shaped half-rings and their average
length was 3.39 ± 0.12 mm. Both the LLD and DVD of
these rings decreased caudally. The distance between the
medial ends of the rings also decreased caudally from
3.56 ± 0.17 mm to 2.74 ± 0.16 mm and this distance was
covered by MTM (Fig. 1).
The medial walls of the right and left bronchus
primarius fused at the level of the bifurcatio tracheae
and formed the pessulus, which extended dorsoventrally
as a double-folded mucous membrane (average
dorsoventral length 4.00 ± 0.17 mm). A ligamentum
interbronchiale connecting the right and left bronchus
primarius was not observed (Fig. 1). Whereas the
intrinsic muscles were lacking in the syrinx, there were
two extrinsic muscles. The sternotracheal (ST) muscle,
one of the extrinsic muscles of the syrinx, was located
(average 14.82 ±.0.36 mm) above pessulus. The
tracheolateral (TL) muscle, another extrinsic muscle was
located at the side of the trachea laterally of the ST
muscle to MTL and it was inserted on MTL (Fig. 2).
In histological sections, taken from the level of
the third and fourth CTS, the lamina epithelialis of the
mucosa was lined with non-cornified stratified
squamous epithelium. There were cilia on the apical
surfaces of the epithelial cells. The lamina propria and
the submucosa which were placed under the epithelium
were composed of compact elastic fibres and loose
connective tissue, respectively. The most peripheral
portion of the propria-submucosa of lamina epithelialis
contained small numbers of mucous glands. Pincus acid
orcein-Giemsa stain revealed that the submucosa
displayed dark-brown stained elastic fibers. Deep to this
loose connective tissue was a prominent stratum of
longitudinally oriented elastic fibers between the hyaline
cartilages of the syrinx. A longitudinal muscular layer
was found throughout the section. The outermost layer
of the section, tunica adventitia, consisted of loose
connective tissue. In the trachea, the lamina epithelialis
of the mucosa was lined with pseudostratified,
columnar, ciliated epithelium with goblet cells. The
lamina propria and the submucosa, which contained
loose connective tissue and mucous glands, were found
below the epithelium. Throughout the section, a thin
longitudinal muscle layer was observed in the trachea
(Fig. 3).
Discussion
The morphological structure of the syrinx has
been described in many bird species (4, 8, 9, 11, 15, 22,
24). This study presents some characteristics of this
organ in pigeon. Since both the trachea and bronchus
primarius participated in its formation, the syrinx in the
pigeon could be classified as of tracheobronchial type
comparable to that of chickens (13, 15, 19). This type of
syrinx is the most common type in birds (13). The
topographical findings of the syrinx in the pigeon were
comparable to those of the chicken (6, 7, 15, 19), new
world pigeon (4)
and ostrich (25).
The tympanum was composed of 5 tracheal
cartilage rings different from those described in the
chicken (2, 6, 7, 15, 18, 19), new world pigeon (4),
singing birds (21), and duck (22). The CTS are
composed of four C-shaped cartilage rings as in
chickens (6, 7, 15, 19) and in ostrich
(25). Although the
CTS was described for several species as joined part of
the tympanum
(15), we found out that in Bursa Roller
Pigeons CTS appeared as a separate cartilage groups.
The CBS were composed of 5 C-shaped half-rings as
described for new world pigeons (4) but different from
those in other birds (15, 18, 19, 22, 24, 25).
The pessulus in pigeons was composed of
connective tissue as described in new world pigeons (4)
and ostrich (25). This structure is cartilaginous in
chickens (6, 15, 18, 19) and ossified in singing birds (11,
24). The connective tissue structure of the pessulus in
pigeons is probably due to the fact that the pigeon is not
a singing bird and thus neither osseous nor cartilaginous
structure is needed to keep the pessulus tight.
Vibrating membranes are present in the walls of
the bronchi. The MTL in pigeons was in the same
location as described for the MTL of new world pigeons
(4) and pigeons (8, 18) and a different location was
found out in the chicken (2, 7, 15, 19), gull (15), singing
birds (24) and ostrich (25). This membrane is defined as
the sound organ in many avian species (15) and the TL
muscle inserts to this membrane so it can produce
significant sound. The MTM covers the open ends of the
CBS as reported in some other species (2, 4, 6, 8, 9, 10,
11, 15, 19, 23).
The syringeal muscles are paired, lie to the
right and left sides of and are divided into two groups,
extrinsic and intrinsic. Extrinsic muscles are the ST and
TL in domestic fowl. Intrinsic muscles are found in
singing birds, suboscine passeriformes, and parrots (2,
16, 21, 23). The TL muscle is found only in the pigeon
(18). In this study, extrinsic muscles, TL and ST, were
observed in pigeon syrinx. The TL muscle was
connected to the MTL, so it might vibrate against MTL
and thus produce the sound. A second muscle, ST,
draws the cartilages to each other and so MTL is kept
under tension.
The histological structure of the syrinx has been
investigated in only few bird species. Of those, the
327
histological structure of the syrinx in the pigeon was
similar to that of the chicken (12), duck (22), and ostrich
(25). The mucosa lining, in particular, was the same as
that of the chicken and duck (3, 5, 12, 22).
This study presents some of the characteristics
of the sound organ in pigeons. Although the
topographical and histological characteristics of the
syrinx showed close resemblance to that of other bird
species, some differences were observed regarding its
anatomy.
References
1. Bacha W.J.: Color Atlas of Veterinary Histology.
Philadelphia, Lippincot Williams & Wilkins, 1990, p.
187.
2. Bell D.J.: Physiology and Biochemistry of the Domestic
Fowl. London, Academic Press, 1971, pp. 100-101.
3. Benks W.J.: Applied Veterinary Histology. London,
Mosby, 1993, p. 404.
4. Bettina M., Pablo L.T.: Relationship between song
characters and morphology in new world pigeons. Bio J
Linn Soc 2001, 74, 533-539.
5. Campbell T.W.: Avian Hematology and Cytology. Iowa,
Iowa State University Press, 1995, p. 55.
6. Dyce K.M., Sack W.O., Wensing C.J.G.: Textbook of
Veterinary Anatomy. London, WB Saunders Company,
1987, p. 785.
7. Getty R.: The Anatomy of the Domestic Animals.
London, WB Saunders Company, 1975, pp. 1899-1902.
8. Goller F., Larsen O.N.: In situ biomechanics of the syrinx
and sound generation in pigeons. J Exp Biol 1997, 200,
2165-76.
9. Goller F., Larsen O.N.: A new mechanism of sound
generation in songbirds. Proc Natl Acad Sci 1997, 23,
14787-14791.
10. Goller F., Larsen O.N.: New perspectives on mechanisms
of sound generation in songbirds. J Comp Physiol A
Neuroethol Sens Neural Behav Physiol 2002, 188, 841-
850.
11. Hartley R.S., Suthers R.A.: Lateralization of syringeal
function during song production in the canary. J
Neurobiol 1990, 21, 1236-1248.
12. Hodges R.D.: The Histology of the Fowl. London,
Academic Press, 1974, pp. 128-131.
13. Hummel G.: Anatomie und Physiologie der Vögel.
Stuttgart, Verlag Eugen Ulmer GmbH, 2000, pp. 116-
118.
14. International Committee on Avian Anatomical
Nomenclature. Handbook of Avian Anatomy: Nomina
Anatomica Avium, Cambridge, The Nuttall
Ornithological Club, 1993, pp. 257-299.
15. King A.S.: Functional anatomy of the syrinx. In: Form
and Function in Birds, edit by King AS, McLelland J.
London, Academic Press, 1989, pp. 105-182.
16. König H.E.: Anatomie und Propädeutik des Geflügels.
Stuttgart, Schattaver, 2001, p. 110.
17. Luna L.G.: Manual of Histologic Staining Methods of the
Armed Forces Institue of Pathology. McGraw-Hill Book
Company, 1968, p. 77.
18. McLelland J.: Avian Anatomy. England, Wolfe
Publishing Ltd, 1990, p. 100.
19. Nickel R., Schummer A., Seiferle E.: Anatomy of the
Domestic Birds. Berlin, Verlag Paul Parey, 1977, p. 65.
20. Osmanlı A.K.: Çeşitli güvercinlerin cins ve özelliklerini
bildirir bir pusula. Istanbul 1923, 40, 18-19.
21. Robinson M.C.: Laboratory Anatomy of the Domestic
Chicken. Iowa, Brown Company Publishers, 1970, p. 8.
22. Scala G., Corona M., Pelagalli G.V.: The structure of the
syrinx in the duck (Anas platyrhinchos). Anat Histol
Embryol 1990, 19, 135-142.
23. Seller T.J.: Bird Respiration. Florida, CRC Press, 1987,
pp. 72-75.
24. Warner R.W.: The anatomy of the syrinx in passerine
birds. J Zool 1972, 168, 381-393.
25. Yıldız H., Bahadır A., Akkoç A.: A study on the
morphological structure of syrinx in ostriches (Struthio
camelus). Anat Histol Embyrol 2003, 32, 187-191.
... These cartilages combine forming the tympanum. It was reported that 4 trachea rings form the tympanum in Denizli roosters, 3 in red hawks; 3 in ostriches, and 2 in geese (Kabak et al., 2007;Tasbas et al., 1994;Yıldız et al., 2005). ...
... Pessulus is located at the end of the tympanum, consist of different structures that vary among species. It consists cartilage in chickens and quails, connective tissue in species such as pigeons (Bayram and Liman, 2000;Getty, 1975;King and Mclelland, 1984;Nickel et al., 1977;Yıldız et al., 2005). The pessulus does not exist in penguins, pelicans and loons (Griffiths, 1994;Tasbas et al., 1994). ...
... In geese (Onuk et al., 2010), pigeons (Yıldız et al., 2005), seagulls (Gezer et al., 2012), ostriches (Yıldız et al., 2003), ducks (Frank et al., 2007) and in some domestic birds tracheobronchial-type syrinx was reported (Bayram and Liman, 2000;Cevik-Demirkan et al., 2007;Kabak et al., 2007;King, 1989). Similarly, in this study budgerigars and canaries presented to have tracheobronchial-type syrinx. ...
Macroanatomic and Histological Examination of the Trachea and Syrinx in Budgerigars and Canaries
Article
Full-text available
  • Jun 2020
The purpose of the present study is to examine and compare the trachea and larynx caudalis of budgerigars and canaries in a macroanatomic manner. 18 canaries and 29 budgerigars that were collected as dead from the clinics of the surrounding area were used in this study. The number of cartilago trachea was 46-47 in canaries; and 53-61 in budgerigars. Crossman's Triple Staining Method was used for histological examinations. 0.1% methylene blue solution was used to examine in a stereo microscope. Tracheobronchial-type syrinx was observed in both species. In canaries last 4-5 trachea rings were joined together to form the tympanum. It was observed that in all species the trachea rings were full. The ligamentum annulare and musculus trachealis were not observed in both species. While pessulus and tympanum were observed in canaries, they were not detected in budgerigars. Musculi syringeales was detected in budgerigars in an extremely developed structure. Cartilagines bronchosyringeales formed the last part of the syrinx; and it consisted 8-9 cartilages in canaries and 6-7 cartilages in budgerigars. Özet: Bu çalışmanın amacı, muhabbet kuşları ve kanaryaların trachea ve larynx caudalis'ini karşılaştırmalı olarak makroanatomik incelemektir. Çevre kliniklerden ölü olarak toplanan 18 kanarya ve 29 muhabbet kuşu bu çalışmada kullanıldı. Cartilago trachealis sayısının kanaryalarda 46-47, muhabbet kuşlarında ise 53-61 olduğu görüldü. Histolojik incelemeler için Crossman üçlü boyama yöntemi kullanılmıştır. Stereo Mikroskopta incelemek içinse % 0.1 metilen mavisi çözeltisi kullanılmıştır. Her iki türde de tracheobronchial tipte syrinx gözlendi. Trachea kıkırdaklarının tam bir halka şeklinde olduğu gözlendi. Ligamentum anulare ve musculus trachealis iki türde de görülmedi. Kanaryalarda pessulus ve tympanum görülürken, bu yapılara muhabbet kuşunda rastlanılmadı. Muhabbet kuşlarında ise oldukça gelişmiş yapıda musculi syringeales görüldü. Cartilagines bronchosyringeales'in syrinx'in son kısmını oluşturduğu görüldü; bu kısmı kanaryalarda 8-9 kıkırdak, muhabbet kuşlarında ise 6-7 kıkırdak meydana getirmişti.
... It is called the tracheobronchial type because both trachea and primary bronchi participate in its formation. The tracheobronchial syrinx is found, for example, in chickens (King, 1989), turkeys (Ragab et al., 2016), pigeons (Yildiz et al., 2005), quails (Cevik-Demirkan et al., 2007), mallards (Yilmaz et al., 2012), geese (Onuk et al., 2010), ostriches (Yildiz et al., 2003), Denizli roosters (Taşbaş et al., 1994), and grey parrots (Warren et al., 1996). However, there is a wide variation in the structural and morphological characteristics of syringes in all these bird species. ...
... The morphological features of the syrinx have been described in several bird species by many authors (Myers, 1917;Nottebohm, 1976;Yildiz et al., 2003Yildiz et al., , 2005Frank et al., 2006;Khaksar et al., 2012;Keskin & Ili, 2013;Erdogan et al., 2015;Ozudogru et al., 2015;Al-aameli & Kadhim, 2017). However, these investigations did not constitute a coherent presentation for cellular organization and ultrastructural features of the syringes in different bird species. ...
... The syrinx of fowls and pigeons could be classified as tracheobronchial in type as described by Cover (1953), Nickel et al. (1977), and King (1989) in fowls and by Yildiz et al. (2005) in pigeons. The present results revealed that in fowl and pigeon, syrinx was formed of three main parts including tympanum (cranial) part, intermediate syringeal part, and bronchosyringeal (caudal) part, in addition to pessulus and tympaniform membranes. ...
Comparative Morphological Features of Syrinx in Male Domestic Fowl Gallus gallus domesticus and Male Domestic Pigeon Columba livia domestica : A Histochemical, Ultrastructural, Scanning Electron Microscopic and Morphometrical Study
Article
Full-text available
  • Jan 2020
Many studies have been carried out to investigate the morphological structure of the syrinx in many bird species. However, the cellular organization of the syrinx in the fowls and pigeons is still unclear. The current study revealed that in fowl and pigeon, the syrinx is formed of three main parts including tympanum (cranial) part, intermediate syringeal part, and bronchosyringeal (caudal) part, in addition to pessulus and tympaniform membranes. A great variation in the structural characteristics of syrinx of fowl and pigeon was recorded. In fowl, the tympaniform membranes showed a characteristic distribution of elastic and collagen fibers which increase the elasticity of tympaniform membranes. Moreover, the bony pessulus helps the medial tympaniform membranes to be stiffer, vibrate more strongly so that louder sound will be generated. In pigeon, the lateral tympaniform membrane is of greater thickness so that the oscillation of this membrane is reduced and the amplitude is lower. Moreover, the pessulus is smaller in size and is formed mainly of connective tissue core (devoid of cartilaginous or bony plates), resulting in the failure of stretching and vibrating of the medial tympaniform membranes, that leads to the generation of deeper sound. Electron microscopic examination of the syringes of fowls and pigeons revealed numerous immune cells including dendritic cells, plasma cells, mast cells, and lymphocytes distributed within syringeal mucosa and invading the syringeal epithelium. Telocytes were first recorded in the syrinx of fowls and pigeons in this study. They presented two long telopodes that made up frequent close contacts with other neighboring telocytes, immune cells, and blood capillaries.
... The topographical findings of the syrinx in male adult fowl in this study were similar to those of mallards [8,9] geese [ 10,11] Denizli roosters [12] pigeons [13,14] turkeys [15][16][17] guinea fowls [18] and ostriches [19]. It could be classified as tracheobronchial in type as described in most common birds such as pigeons [14] quails [20,21] mallards [8,9,22] geese [10,11] ostriches [19] long legged buzzards [23] Denizli roosters [12] guinea fowls [18] black francolins [24] and eagle owls [21]. ...
... The topographical findings of the syrinx in male adult fowl in this study were similar to those of mallards [8,9] geese [ 10,11] Denizli roosters [12] pigeons [13,14] turkeys [15][16][17] guinea fowls [18] and ostriches [19]. It could be classified as tracheobronchial in type as described in most common birds such as pigeons [14] quails [20,21] mallards [8,9,22] geese [10,11] ostriches [19] long legged buzzards [23] Denizli roosters [12] guinea fowls [18] black francolins [24] and eagle owls [21]. ...
... The finding obtained from this investigation was in agreement with Myers and Meclelland [26,27] and dissimilar from the result of Freeman Dyce et al., and King [28][29][30] which recorded that the tympanum was consisted of three or five tracheal rings in chickens. While the number of tracheal rings forming the tympanum were reported as two in quails [20,21], three in long legged buzzards [23] ostriches [19] guinea fowls [18] turkeys [31] and song birds [32] five in sea gulls [33] and pigeons [14]. ...
The Gross Anatomy of the Syrinx of Adult Male Domestic Fowl Gallus gallus domesticus
Article
Full-text available
  • Aug 2019
Birds are considered a good model for studying the phonation process, the syrinx is a vocal organ in birds. The purpose of this study is to investigate the topographical and morphological characteristics of syrinx of male domestic fowl. In the current study we use the syringes of seven adult males. The study shows that the syrinx of investigated birds is tracheobronchial in type. It consists of; tympanum, tracheosyringeal and bronchosyringeal groups. In addition, there are interbronchial ligament (brachidesm), lateral and medial vibrating membranes as well as the pessulus at the tracheal bifurcation. Tympanum part forms the first part of the syrinx; it is formed of four tracheal rings. The tracheosyringeal part is located at the point of tracheal bifurcation just below the tympanum. It is formed of four highly modified incomplete tracheal rings. The bronchosyringeal part is formed of first three pairs of bronchial half-rings. The current study was presented the detailed morphological characteristics of syrinx in male domestic
... The tracheobronchial type was commonly collected in ostrich syrinx (Yildiz et al., 2003), Bursa Roler Pigeons (Yildiz et al. 2005), white turkey (Arican et al., 2007 andKhaksar et al., 2012), long-legged buzzard (Kabak et al., 2007) and quails (Bayram andLiman, 2000 andÇevik et al. 2007), Greater Rhea (Rhea Americana) (Picasso and Carril, 2013), in guinea fowl (Al-Bishtue, 2014), sparrow hawk (Ozudogru et al., 2015), baladi turkey (Ragab et al., 2016), black francolin (Al-Aameli and Kadhim, 2017), white pekin ducks (Mohamed R., 2017). ...
... Anatomical structure of syrinx has been investigated in many bird's species such as falconid (Griffiths, 1996), Ostrich syrinx (Yildiz et al., 2003), Bursa Roler Pigeons (Yildiz et al. 2005), Greater Rhea (Rhea Americana) (Picasso and Carril, 2013), sparrow hawk (Ozudogru et al., 2015), baladi Turkey (Ragab et al., 2016), chukar partridge (Galliforms) (Erdoğan et al., 2015), Sparrow hawk (Ozudogru et al., 2015), black francolin (Al-Aameli and Kadhim, 2017), White Pekin Duck (Mohamed R., 2017). The tympanum constituted of the cranial cartilages only (Khaksar et al. 2012) or the tympanum compromised both the cranial and the intermediated cartilages (O'Malley, 2005). ...
... The type of the lesser kestrel syrinx is tracheabronchial, which is similar to the observations on many avian species; in ostrich syrinx (Yildiz et al., 2003), Bursa Roler Pigeons (Yildiz et al. 2005), white turkey (Arican et al., 2007 andKhaksar et al., 2012), long-legged buzzard (Kabak et al., 2007) and quails (Bayram andLiman, 2000 andÇevik et al. 2007), Greater Rhea (Rhea Americana) (Picasso and Carril, 2013), in guinea fowl (Al-Bishtue, 2014), sparrow hawk (Ozudogru et al., 2015), baladi turkey (Ragab et al., 2016), black francolin (Al-Aameli and Kadhim, 2017), white pekin ducks (Mohamed, 2017). ...
Anatomical study on the syrinx of the lesser kestrel (Falco naumanni)
Article
Full-text available
  • Jan 2018
Infection of falcon by aspergillus fumigatus, asp Flavus and asp Niger cause change in the voice, inability to vocalize or respiratory noises, due to the syrinx lesions, also severe dyspnea occur when obstruction happened. Due to the little available literatures about falcon anatomy, we have done this study to investigate the structure of the lesser kestrel syrinx as it is the voice box in birds which differ from a species to the other. Lesser kestrel (Falco naumanni) syrinx is tracheobronchial in type. The tympanum is composed of the last three cartilaginous tracheal simple complete rings. The tracheosyringeal group is formed of four fused single, ossified, special, characteristic rings just before tracheal bifurcation. The bronchosyringeal group is consisted of three paired cartilaginous C-shaped rings on both sides of the tracheal bifurcation; the edges of both first rings are single while the edges of the rest two bronchosyringeal rings are double. The first two bronchosyringeal rings are related medially to the medial tympanic membrane while the last one attached to the interbronchial ligament. The pessulus is a stretched triangle in shape, cartilaginous bar in the median plane of the trachea. The lateral and medial tympanic membranes are complete sheathes closing the trachea bronchial junction of syrinx laterally and medially. The lateral one is larger than the medial one, and both membranes are responsible for the sound production.
... The latter foramen was termed by Warner (1971) as the subpessular air space. Moreover, interbronchial ligament connecting the primary bronchi in the present study was similar to many birds, while Yildiz et al. (2005) reported that this ligament is absent in Bursa roller pigeon. ...
... It was recorded in our study that the pessulus in geese was composed of ossified tissue; the same findings were also in guinea fowl (Al-Bishtue, 2014), indigenous male turkey (AL-Mussawy, 2011), avian (Baumel et al., 1993), white pekin ducks (Mohamed, 2017), goose (Onuk et al., 2010), mallard (Yilmaz et al., 2012) and songbirds(Warner, 1972). However(Baumel et al., 1993) in oscine andYildiz et al. (2005) in pigeon stated that the pessulus is formed by a double-folded mucous membrane. On the other hand the pessulus is absent in penguins(Tasbas et al., 1986).The obtained results were parallel to those described in in goose (Onuk et al., 2010) that the bronchosyringeal cartilages were 6 in number, while the bronchosyringeal cartilages are 3 in guinea fowl and francolin (Al-Bishtue, 2014; Kadhim et al., 2017), 4 in turkey (Kookhdan et al., 2012), 5 in pigeon (Yildizet al., 2005) and 7 in sea gulls (Ince et al., 2012). ...
CORRESPONDENCE: Gross anatomical syringeal structures of goose (Anser anser domesticus) http://bdvets.org/javar
Article
Full-text available
  • Dec 2017
Objective: The main purpose of this study was to give detailed information on the position and normal anatomical syringeal structure in goose which had received a little attention in the field of veterinary anatomy. Materials and methods: Six (3 females and 3 males) adult geese weighing 2-4 Kg were used. The goose was slaughtered and its body cavity was opened to detect in situ position of the syrinx. Then the syrinx were dissected and fixed in 10% formaldehyde for 48 h and then kept in 70% ethanol for 2 h. Results: Anatomical examination showed that the syringes of these birds were located in the thoracic cavity at the bifurcation of the trachea. The syrinx was tracheobronchial type formed by tracheosyringeal cartilages, bronchosyringeal cartilages, pessulus, medial and lateral tympaniform membranes, interbronchial ligament and foramen as well as extrinsic syringeal muscles. Conclusion: There were some similarities and some differences of the anatomical structures of the syrinx of goose and that of other bird species. No differences between male and female syrinx were observed. [J Adv Vet Anim Res 2017; 4(4.000): 343-347]
... The histological features of the syrinx have been described in several bird species by many authors (Yildiz et al., 2003(Yildiz et al., , 2005Frank et al., 2007;Yilmaz et al., 2012;Keskin & Ili, 2013;Ibrahim et al., 2020). However, the cellular elements and the fine structure of the budgerigar syrinx are not fully understood. ...
Anatomical, Histological, and Electron Microscopic Structures of Syrinx in Male Budgerigars (Melopsittacus undulatus)
Article
The syrinx is the main source for phonation in birds, its function is analogous to the mammalian larynx. Birds have both a larynx and a syrinx, but they use only the latter to vocalize. The objective of this work to give a detailed description of the anatomical, histological, and ultrastructural of syrinx in male budgerigars as a model of a passerine bird. The syrinx in the current study was to be found as a tracheobronchial type, it consists of cranial (tympanum) part and caudal (bronchosyringeal) part and, additionally, there are lateral vibrating membranes. The tympanum is formed of the last six tracheal rings, histologically its lamina epithelialis is a pseudostratified ciliated columnar epithelium with goblet cells and interrupted by intraepithelial glands. The secretory acini appear oval and lined by pyramidal secretory cells. The lamina propria–submucosa contain numerous blood capillaries, immune cells, and telocytes (TCs). The electron microscopic examination revealed numerous blood capillaries surrounded by fibroblasts and numerous immune cells, including mast cells and wandering leukocytes, within the tympanum mucosa. Hence, this study provides a detailed knowledge about the syrinx in male budgerigars.
... Similarly, the Yildiz et al. (2003) reported the same findings in ostriches, Yildiz et al. (2005) in pigeons, Frank et al. (2007) in mallards and Khaksar et al. (2012) in female and male turkeys. The tympanum had six complete rings similar to Bursa roller pigeons (Yıldız et al., 2005), long-legged buzzards (Kabak et al., 2007), sea gulls (İnce et al, 2012), geesse (Onuk et al., 2010) and Japanese quails (Çevik et al., 2007). This finding was in contrary to five in pigeons (Yildiz et al. 2005) Table.2 ...
... The present study showed that , pessulus of duck cartilaginous ossified that agreeed with (27) in European starling , (28) in white eared bulbul and, (41) in singing birds, while cartilaginous in chickens (42). While it disagreed with (33,43) , they showed that the pessulus contains connective tissue same as in ostrich and turkeys. ...
COMPARATIVE ANATOMICAL , HISTOLOGICAL AND HISTOCHEMICAL STUDY OF (LARYNX, TRACHEA AND SYRINX) BETWEEN MATURE AND IMMATURE MALES OF LOCAL DUCK (Anas Platyrhnchos )
Article
Full-text available
  • Jan 2020
The present study aims to compare gross anatomical, histological and histochemical structures of the larynx, trachea and syrinx between mature(Drake) and immature (Duckling) male duck(Anas platyrinchous). Twelve healthy local males ducks were divided into two groups. Six Drake at one year old and six Duckling at three months .For anatomical study the larynx, trachea and syrinx were measured by using vernia. For histological study specimens from larynx, trachea and syrinx were fixed in 10% formaldehyde after that the specimens embedded in paraffin. The next step tissues were sectioned using microtome and stained with routine stain and special stains .The anatomical results revealed that larynx, syrinx and trachea are identical in position in both Drake and Duckling , but differ in size. The histological results revealed that larynx of both Drake and Duckling lined by keratinized pseudostratified squamous epithelial tissue that appears thicker in Drake than Duckling. Trachea of Drake and Duckling lined with ciliated pseudostratifed columnar epithelial with mucous glands , supported by ossified cartilages in Drake and hyaline cartilage in Duckling. Syrinx of both Drake and Duckling lining by the ciliated pseudostratifed columnar epithelial with few goblet cells. Beneath it the lamina properia and sub mucosa that contained in loose connective tissue with blood vessels and amount of elastic and collagen fibers beneath it in Drake spongy bone trabeculae that contains bone lacuna fills with osteocytes , while in Duckling hyaline cartilage. The hyaline cartilage consists of lacuna which contains chondrocytes.
... Sirinkste ses oluşumu ile ilgili lateral ve mediyal konumlu iki çift timpanik membran birbirinden bağımsız çalışır ve sirinksi çevreleyen ekstrinsik ve intrinsik kasların gergin halde olmasını sağlar. 5,20 Bu kaslar kıkırdak halkaları farklı derecelerde hareket ettirerek, ses oluşumunda etkin olan membranların gerginliğini değiştirir; böylece sesin tonu ve ritmi de ayarlanmış olur. Denizli tavuğunda LTM ve MTM'na ait lamina propriada daha geniş bir alanda gevşek tarzda yerleşim gösteren elastik fibrillerin submukozaya doğru dalgalı bir görünüm oluşturdukları, erkek sülünde ise elastik fibrillerin geniş bir alanda çok sıkı biçimde düzenlendikleri; bronşiyal kısma ait lamina propriada ise dar bir alanda sıkı biçimde yerleşim gösterdikleri gözlendi. ...
In situ biomechanics of the syrinx and sound generation in pigeons
Article
Full-text available
  • Sep 1997
The in situ biomechanics of the vocal organ, the syrinx, was studied in anesthetized pigeons using fiberoptic instruments. The role of syringeal muscles was determined by electrical stimulation, and phonation was induced by injecting gas into the subsyringeal air sacs. This study presents the first direct observations of the biomechanical processes that occur in an intact syrinx. Contraction of one of the syringeal muscles, the m. tracheolateralis (TL), withdraws the lateral tympaniform membranes (LTM) from the syringeal lumen, causing opening of the syringeal airways. Shortening of a second muscle, the sternotrachealis (ST), draws the syringeal cartilages closer to each other, causing the LTM to fold into the syringeal lumen. Maximal ST contraction does not lead to complete closure of the syrinx. As air-sac pressure is increased by the injection of gas, the LTM are drawn into the syringeal lumen and balloon in a rostral direction until they touch, thus forming a fold-like valve. Air-induced phonation is always associated with vibrations of the membrane folds, suggesting that pulsatile release of air into the trachea by vibratory motion of the LTM generates sound. During air-induced phonation, strong stimulation of the TL terminates sound generation by abducting the LTM, whereas weak stimulation changes the geometry of the membrane folds, which is accompanied by changes in the acoustic structure of the sound. Stimulation of the ST has little effect on air-induced sounds. The LTM appear to be the main sound generators, since disabling the medial tympaniform membranes (MTM) with tissue adhesive does not prevent phonation or change the frequency and amplitude structure of display coos in spontaneously vocalizing pigeons. Moreover, the activity of the syringeal muscles appears to have a mainly modulatory function, suggesting that the basic sound-generating mechanism is similar in both air-induced and natural phonation.
A new mechanism of sound generation in songbirds
Article
Full-text available
  • Jan 1998
Our current understanding of the sound-generating mechanism in the songbird vocal organ, the syrinx, is based on indirect evidence and theoretical treatments. The classical avian model of sound production postulates that the medial tympaniform membranes (MTM) are the principal sound generators. We tested the role of the MTM in sound generation and studied the songbird syrinx more directly by filming it endoscopically. After we surgically incapacitated the MTM as a vibratory source, zebra finches and cardinals were not only able to vocalize, but sang nearly normal song. This result shows clearly that the MTM are not the principal sound source. The endoscopic images of the intact songbird syrinx during spontaneous and brain stimulation-induced vocalizations illustrate the dynamics of syringeal reconfiguration before phonation and suggest a different model for sound production. Phonation is initiated by rostrad movement and stretching of the syrinx. At the same time, the syrinx is closed through movement of two soft tissue masses, the medial and lateral labia, into the bronchial lumen. Sound production always is accompanied by vibratory motions of both labia, indicating that these vibrations may be the sound source. However, because of the low temporal resolution of the imaging system, the frequency and phase of labial vibrations could not be assessed in relation to that of the generated sound. Nevertheless, in contrast to the previous model, these observations show that both labia contribute to aperture control and strongly suggest that they play an important role as principal sound generators.
Relationship between song characters and morphology in New World pigeons
Article
  • Dec 2001
We studied the pattern of variation in song characters among 16 New World pigeon species belonging to different taxonomic groups defined by morphological characters. Structural, temporal and frequency characters of the song were analysed. Principal components analyses showed that species belonging to the same taxonomic group were also grouped together by their song characters. In addition, individuals were correctly assigned into taxonomic groups by discriminant function analyses in more than 87.8% of cases. These analyses also showed that more than 87.5% of the individuals could be correctly classified by species when all song characters were included. Correct classification of individuals by species and taxonomic groups dropped when character types were analysed separately, thus showing that structural, as well as temporal and frequency characters are fundamental to define species- and group-specific identities of New World pigeon's songs. Correspondence between patterns of vocal and morphological variation found in this study can be a consequence of evolutionary changes in morphology affecting song production, as for example body size changes that constrain the syrinx to produce certain acoustic frequencies.
[The structure of the syrinx in the duck (Anas platyrhinchos)]
Article
  • Jul 1990
The morphological and structural aspects of the Membrana tympaniformis medialis of the syrinx were studied with L.M. and E.M. in 32 female and male Peking ducks from 6 to 12 months old. The membrane is covered by pseudostratified prismatic epithelium which shows different regional features. While at level of the proximal and distal third of the tympanic membrane the morphological and structural aspects of the cells resemble that of the adjacent structures (pessulus and primary bronchi), the middle part of the epithelium layer becomes gradually flattered; the epithelium is made up of oblique or horizontal cells which touch each other by means of many lateral interdigitations. At level of the upper parts of the cells there are several types of cell junctions. This particular disposition seem to be well adapted to the contractions of the syringeal muscles.
Lateralization of syringeal function during song production in the canary
Article
  • Dec 1990
The canary (Serinus canaria) vocal organ, the syrinx, has two separate sound sources, one in the cranial end of each bronchus. Previous investigations of whether song syllables are produced unilaterally or bilaterally have provided two contradictory results, as one researcher suggested that almost all syllables are produced by the left side of the syrinx alone, whereas another researcher suggested that both sides contribute similarly to all syllables. Our experiments, which involved unilateral bronchus plugging followed later by denervation of the ipsilateral syringeal muscles, attempted to resolve this disagreement. The males with right bronchus plugs, singing on the left side of the syrinx alone, produced nearly normal songs, whereas the birds with left bronchus plugs, singing on the right side, sang quite poorly. Interpretation of these data is difficult because it is not clear how syringeal function would be affected if the airflow rate through the intact side is increased above normal, nor is it known if the bird can compensate for bronchus occlusion. Nonetheless, we suggest that in male canaries most syllables are normally sung by the left side alone, with some syllables being produced by the right side alone and some being sung by both sides together. Right nerve section had little effect on the right-bronchus-plugged males' ability to sing, but the repertoires of the left-plugged males were altered after left nerve section, indicating the possibility that signals carried by the left nerve exert an influence on the contralateral side.
New perspectives on mechanisms of sound generation in songbirds
Article
  • Jan 2003
The physical mechanisms of sound generation in the vocal organ, the syrinx, of songbirds have been investigated mostly with indirect methods. Recent direct endoscopic observation identified vibrations of the labia as the principal sound source. This model suggests sound generation in a pulse-tone mechanism similar to human phonation with the labia forming a pneumatic valve. The classical avian model proposed that vibrations of the thin medial tympaniform membranes are the primary sound generating mechanism. As a direct test of these two hypotheses we ablated the medial tympaniform membranes in two species (cardinal and zebra finch) and found that both were still able to phonate and sing without functional membranes. Small changes in song structure (harmonic emphasis, frequency control) occurred after medial tympaniform membrane ablation and suggest that the medial tympaniform membranes play a role in adjusting tension on the labia. Such a role is consistent with the fact that the medial tympaniform membranes are directly attached to the medial labia. There is no experimental support for a third hypothesis, proposing an aerodynamic model for generation of tonal sounds. Indirect tests (song in heliox atmosphere) as well as direct (labial vibration during tonal sound) measurements of syringeal vibrations support a vibration-based sound-generating mechanism even for tonal sounds.