R. Nickel -A.Schummer - E. Seiferle Anatomy of the Domestic Birds Translation by W.G. Siller and PA.L.Wight With 141 illustrations, some in colour, in the text and on 7 plates 1977 Verlag Paul Parey Berlin - Hamburg | Mi30 Muscles of the trunk to the skull in the bird and is therefore very mobile The m. stylohyoideus (29/6) arses laterally fon che angular bone of the lower jaw and isin two parts, one of which is inserted to the basihyoid and the other to the urohyale. Its function isto lift and retract the hyoid bone and the cranial larynx. The m. keratoglossus (/7) arises from the rostral part of the ramus of the hycid bone and is inserted on the lateral border of the entoglossal bone, Bilateral traction pulls che vongue downwards, unilateral traction pulls it to one side. The m. keratohyoideus (/8) arises from the inner side of the ramus of the hyoid bone and is inserced on the urobyale. Ies action is to bring about apposition of the rami of the hyoid bone. ‘The m. hypoglossis obliquus arises on the undersurface of the basihyale and ends on the lateral border of the entoglosal bone. The tip of the tongue is raised by the action of this muscle. ‘The m. hypoglossus rectus arises ventrally on the basihyale and ends on the ventral surface of the entoglosal bone. This muscle pulls down the tongue. The m. thyreohyoideus arises onthe lateral sucface of the thyroid cartilage and is inserted ‘on the basihyale and the entoglostal bone. It represents the cranial elongation of the m. sterno- thyreoideus (/9; 31/47) which extends from the lateral surface of the trachea to the thyroid cartilage, The function of both these muscles isto pull down the tongue and the cranial larynx. Muscles of the trunk (29-32) ‘Muscles of the head joints Abbreviations used in this section are as follows: — (©. = origin; I. = insertion; F. = function; N. = nerve supply. M. rectus capitis dors. major surface of the occipital bones the eervical nerves. M. obliquus capitis cran.; O.: Spinous process of the axis; I Occipital bone; F.: To extend the oceipito-atlantal joine and rotate the atlanco-axial joint; N.: Dorsal branches of the cervical M. obliquas capitis caud.s Lateral surface of the atlas; F. cervical nerves. M. rectus capitis vente, lat. (29, 30/11; 31/5); Ou: Transverse processes of the 4th and Sch cervical vertebrae; I.: On the basal surface of the occipital bone; F.: To flex the occipital joints Ventral branches of the cervical necves, 'M. rectus capitis ventr. med. (29/12; 31/6); O.: Bodies of 2nd to 7th cervical vertebrae; Basally on the occipital bone; Faz To flex the occipital joint; N.: Ventral branches of the cervical nerves. M. longus colli (31/7); O.: Pleurapophyses of the last cervical vertebrae; I: The pleurapo- physes of the cranial thoracic vertebrae and cervical ribs up to the 2nd cervical vertebra; F. To flex the neck; N.: Ventral branches of the cervical nerves. Spinous process of cervical vertebrae 1—3; I: Nape 'o extend the occipito-atlantal joints; N.t Dortal branches of orsolateral surface of the 2nd co 4th cervical vertebrae; 1: To rotate the atlanto-axial joint; N.t Dorsal branches of the ‘Muscles of the vertebral column M. iliocostalis; O.: Craniodorsal border of che ilium, transverse processes of the lumbar and sacral vertebrae and the ribs; Lz Cranially it runs into the longissimus dorsi muscles F.: To support the vertebral column and theyribs; N.: Dorsal branches of the thoracic, lumbar and sacral nerves. M. longissimus dorsi; O.: Craniodorsal border of the ilium, transverse processes of the lumbar vertebrae; L.: Spinal and transverse processes of the preceding vertebrae; F.: To support the vertebral column; N.: Dorsal branches of the thoracic, lumbar and sacral nerves. M, longissimus cervicis (31/1); O.: Transverse processes of the Ist to 8th thoracic vertebrae; I: Transverse processes of the last five cervical vertebrae; F.: To extend the cervical vertebral column; Ne: Dorsal branches of the cervical and thoracic nerves.Thoracic and abdominal muscles 31 Iaterally M, semispinalis cervcis(/2); 0.1 "Transverse processes of the Sch to 1th cervical vertebrae; oasihyoid I Anicular proceses of the same cervical vertebrae, 50 that muscle bundles are inserred 20 we cranial alternate into the aticolar process of the next vertebrae but one; F.z To extend and support id bone the cervical vertebral eolumns Nez Dorsal branches of the cervical nerves, ze rongee pinalis capitis consists of: from the 4) M. biventer cerviis (3); Ou: Ac the level of the tse shoracie vertebra from the m.spinalis to bring Nape surface of the occipital bone; Fut To extend the ocepito-atlantal and the aanto- son the zal join and the cervical vertebral column; Nit Dorsal branches of the cervical nerve. Sip of the 1b) M. complexus major (29, 30110; 31/4); Our Transverse processes of the 2nd to Sth cervical ly on the, vertebrae; I.: Occipital bone; F.: To extend the occipito-atlantal and atlanto-axial joints and own the the eervial verebraleolumn; Ne: Dorel branches ofthe cervical nerves s inserted ‘M. spinalis dorsi; Ou: Spinous processes of the thoracic vertebrae and the last cervical vertebrae; tu sterno- 1: Tndevidual bundles are inserted onto the articular processes of the free thoracic and last > thyroid, cervical vertebrae; F.: To extend the thoracic and elevate the cervical vertebral column; N.: rynx. Dorsal branches of the cervical and thoracic nerves. M. spills cervicis; Ou: Spinous procestes ofthe 11th to 2nd cervical vertebrae; Li Transverse procawes of the 3th to foe cervical vertsbrae; Fs To extend and elevate the cervical vertebral Eslumnn; Ne: Dorsal branches of the cervical nerve. ‘ML muliiidus dorsi et cervics; Ou Transverse process of 3rd choracic to deh cervical verubrae; Lz Spinous proceses of each preceding vertebra; Fz To support and elevate the cervical Yertebral column; Nez Dorsal brandies ofthe cervical and thoracic necves ‘Mim. rotatores; These muscles are derived from the m. multifidas and ran beoween the wans- vene and he caudal articular process of che cervical vertebrae; Bz To rotate the verteraes N.: Dorsal branches of the cervical nerves. ‘Mon interopinales; These musles connect neighbouring spinous procestes between the 2nd to S ‘11th cervical vertebrae and they may also extend to the next vertebra but one; F.: To fix and Li Nape ‘move neighbouring vertebrae; Nc: Dorsal branches ofthe cervical nerves. anches of ‘Mm, intertransversarii; These muscles link the transverse processes of successive vertebrae up . to the Bed cervial; Ft To fix the vertebral column and to bend the cervical vertebral column Oona Interally; Net Dossal branches of the cervical and thoraci nerves. ebrae; Lz Muscles of the thoracic wall oe Mm. intercostales ext.; O.: Caudal borders of the ribs and uncinate processes; I: Cranial borders rn ofthe succeeding ribs; Fe Inspiration; Nu intercostal nerves. oe ‘Mim. intercostals intj O.2 Cranial borders ofthe ribs L: Caudal borders of the preceding ribss aaa Fi: Expiration; Nu: Intercostal nerves. a ‘Mim, levatores costarum; O.: Transverse processes of the 2nd to 5th thoracic vertebrae; I rot the ee eC branches of the thoracic nerves. M. scalenus; Ou: Transverse process of the second last cervical vertebra; L: Cranial borders of the ist and 2nd ribs; F.: Auxiliary inspirator; N.z Ventral branches of the last cervical and first shoracie nerve. ™M. subcostalis; scernal ribs; Fr Auxiliary inspirator; raniolateral process of the sternum; I.t Internal surface of the 3rd to 6th intercostal nerves. Muscles of the abdominal wall anbar and ‘ support M. obliquus abdominis ext. (31/8; 32/1); 0: In 4—7 bundles from the laeral surfaces of the al nerves. riby and their uncinate proceses and as an apaneurosis from the iliam and pubis; Lt As a very xe himbar thin aponeurosis on the lateral border of the sternum and into the linea alba; Fa: Abdominal pore the compression and expiration; N.: Intercostal,iliohypogasrie and ilio-inguinal nerves. 'M. obliquus abdominis int. (32/2); O.: Ventral border of the ilium and pre-acetabular border sercebrac; of the pubis: L: caudal border of the last vertebral nb. and union with the aponcuross of the vertebral sm. rectus abilominis: Ft To compress the abdomen and bring aboot expiration; Net Intercorl, iliohypogastric and ilioinguinal nerves.32 ‘Muscles ofthe limb M. rectus abdominis (32/3); O.t Lateral borders of the sternum and caudal border of the last sternal rib; I Ventral border of the caudal half of the pubic bone; F.: To cooperate in abdominal compression and expiration, and to elevate the sternum; N.: Intercostal,iliohypogastrie and ilio- inguinal nerves. 'M. transversus abdominis (/4); O.: Ventral border of the pubis and pre-acetabular part of the slfum, internal surface of the last shree ribs; Ls Internal surface of the sternum and median union of the aponeurosis with that of the opposite side; F.: To contract the abdomen, expiration; N.t Intercosta, iliohypogastric and ilio-inguinal nerves. ™M. levator ani (31/9); O.: Caudal process of the ilium; L: Median union of the two aponeu- soses, caudally it joins the m, sphincter ani; F.:To elevate the anus; Na: Sacral nerves. Muscles of the tail M. levator coceygeus (/10); O.t Caudal border of the ilium and the ischium and the transverse processes of the last lumbar and most of the coceygeal vertebrae; I: Spinous processes of the Coceygeal vertebrae and the pygostyle; F.: To raise the tail and pull it to the side; N.t Sacral and coceygeal nerves. ‘M. coccygeus lat. (/11); Ou: The transverse processes of the free coceygeal vertebrae; Ls Ventral surface of the body of the succeeding coccygeal vertebrae and the pygostyle and also the rectrices fo steering feathers; F.: To pull down the tail and spread the rectrices; Nu: Sacral and coccygeal 'M. coceygeus (/12); O.t Caudal end of the pubis and ischium; I: 2—4 outer rectrices, che transverse processes of the free coceygeal vertebrae and the pygostyle; F.z To pull the tail sideways and downwards and to spread the rectrices; N.+ Sacral and coccygeal nerves. M. iligcoceygeus; O.: Dorsal surface of the ilium and dorsal surface of the coccygeal vertebrae; La 4-8 of the outer recirices; F.: To lift and spread the rectrices; N.: Sacral and coceygeal ‘M. depressor coceygeus; O.: Ventral surface of the last sacral and the free coceygeal vertebracs Lz Ventral surface of the body of the succeeding coccygeal vertebrae and the pygostyle; F.: To ‘pull down the tail; N.: Sacral and coccygeal nerves. Muscles of the limbs (1, 32) Muscles of the pectoral limb ‘Trunk-limb muscles M. trapezius (31/14); Out Spinous processes of the first to sixth thoracic vertebrae; I Vertebral border of the scapula; F.: To lift the scapula; N.t Dorsal cervical and accessory nerves. M. thomboideus supf.; O.: Spinous processes of the last cervical and first choracic vertebrae; L: Proximal part of the farcula; F.t To assis in expiration; N.z Dorsal thoracic nerves of the brachial plexus. M. rhomboideus prof.; ©. Spinous processes of the last cervieal and first to fifth thoracic vertebrae; I: Dorsomedial border of the scapula; F.: To raise the scapula; Nu: Dorsal thoracic nerves of the brachial plexus. ‘M. serratus ventr. supf. consists of three parts: Pars eran,; O.+ First rib; I: Middle third of the ventral border of the scapula; Pars caud, (31/13); O. Outer surface of the uncinate process of the sternal ribs; 1: Last third of the ventral border of the scapula; F.+ To pull down scapula and assist in expiration, Pars metapatagialis; O.: Outer surface of the uncinate process of the sternal ribs; L: Patagium between the trunk and upper arm; F.: To stretch the patagium; N.: Dorsal thoracie nerves of the brachial plexus. ™M. serratus prof. ‘ransverse processes of the last cervical vertebrae as well as the first and second ribs; L: Medial surface of the scapula; F.: To lift and abduct the scapula and assist in expiration; Nut Dorsal thoracic nerves of the brachial plexusof the last abdominal cand ilio- art of the dian union © aponeu- sses of the Sacral and Ls Ventral ‘coveygeal strices, the Il the ail vertebrae coceygeal vertebrae; dle; Fer To + Vertebral vertebrae; vves of the fh thoracic al thoracic Last third + Paragium ves of the ve firse and id assise in ‘Mascles of the pectoral limb 33 1M, latissimus dorsi (/16); 0. In ewo heads from the spinous processes of the two first and the last thoracic vertebrae; I: Crest of the lateral tuberosity of the humerus; F.: To fold up and carry the wing during rest; N.: Branches of the dorsal brachial nerves of the brachial plexus. ™M. pectoralis supf, (51/17; 32/3); O.: Venteal border of che lateral crest and process of the sternom, the clavicle and the sternal ribs; I: Crest of the lateral tuberosity of the humerus; F.: To perform the downstroke of the wings; N.: Pectoral nerve of the brachial plexus. M. pectoralis prof. (32/6); O.: Lateral surface of the sternum and coracoclavicular membranes Iu: Lateral tuberosity and crest of the lateral cuberosity of the humerus; its terminal tendon runs through che foramen triosseum; F.z To life the wing; Net Pectoral nerve of the brachial plexus ‘Muscles of the shoulder and upper arm M, deltoideus major (51/18); O.: Scapula, clavicle and coracoid; L ‘The crest on the lateral tuberosity of the humerus; F.: To flex the shoulder joint; Nu: Axillary nerve, M, deltoideus minor; O.: Scapula and claviele; L Lateral tuberosity of the humerus; F.: To life the wing; N.t Axillary nerve. M. seapulohumeralis cran, O.: External surface of che scapula; I Jhumerus; F.z Adducts and lifts the upper arm; N. Subscapular nerves. ‘M. scapulohumeralis caud. (/19); O.: Ventral border and outer surface of the scapula; It Medial tuberosity of the humerus; F.: To lift the wing; N.: Ventral thoracic nerves of the brachial plexus. 'M. subscapularis; O.: Lower ventral border and costal surface of the scapula; I: Medial tuberosity of the humerus; F.: To perform the downstroke of the wings; Na: Ventral thoracic nerves of the brachial plexus. 'M. biceps brachii (32/7); O.: Coracoid, clavicle and proximal end of the humerus; L: Proximal end of the radius and ulna; F.+ To flex the elbow joint; N.: Medio-ulnar nerve. 'M, brachialis (32/8); O.: Distal end of the humerus, proximal to the medial condyles Lt Proximal end of the ulna; F-: To flex the elbow joint; N.: Medio-ulnar nerve, M. coracobrachialis (/9); O.: Coracoid bone and sternocostal bones; tuberosity of the humerus; F.: To pull down the upper arm; Nu: Axillary and medio-ulnar nerves. M, supracoracoideus; O.: Cranial border and outer surface of the coracoid bone; I Crest on the lateral tuberosity of the humerus, the tendon goes through the foramen triosseum; F.t To lift the wing; N.t Medial thoracic nerves of the brachial plexus. M, sternocaracoideus; O.: Lateral process of the sternum; I Distal end of the coracoid bones Fi: To hold the sternocoracoid articulation; N.1 Medial thoracic nerves of the brachial plexus. 'M. triceps brachii (31/20; 32/10); O.t Caput longum on the neck of the scapula, caput bumerale in ewo parts on the inner surface of the head and on the medial tuberosity of the humerus; Olecranon of the ulna; F.: To extend che elbow joint; Ne: Anconaeus and radial nerves. M. anconaeus; O.: Medial epicondyle of che humerus; I: Proximally on the radial surface of the ulias F.s To extend the elbow joint; N.: Anconaeus nerve. 'M. ectepicondyloulnaris (31/27); O.: Lateral epicondyle of the humeras; L: Radial surface of the nas F.: To extend the elbow joint; N.z Radial nerve. ‘Medial tuberosity of the Muscles of the lower arm and hand M. extensor carpi radials (31/225 32111)s carpale I; Ft Extensor of the carpal join 'M. extensor carpi uinaris (31/23); Ou: Lateral epicondyle of the humerus; Lt Lateral border of, the second metacarpal bones 'M. flexor carpi radials (me. ulnimetacerpalis ventr.) (32/12); O.t Radial and volar susfaces of the ulna; Ls Second metacarpal bone; F: Flexor ofthe carpal joint; N.t Median nerve. ML flexor carpi ulnaris (32/13); O.t Medial epicondyle of the humerus; Li Os carpi ulnare; Fs Flexor of the carpal joint; N.: Ulnar nerve M. ulnimetacarpalis dors. (31/24); Ou: Distal end of the ulnas I Second metacarpal bones F.: To extend the carpal joint; N.: Radial nerveM4 ‘Muscles ofthe limb he surface muscelstore of the fow galerie cop dpreeneand ‘am. complesas sts $m, seme cai ate a 1 longi sevice: 2m G5 ttbittaphis ten meds 7,7 mr lng ells 66m oblige doy 7: PP, pecorae apts Mm deleideay; 9 mapalabarcs ya exempt 9 ms ver psa loge 8m abnor die ice i my tenor lanoe levee wih we imaens nap. (eral part 30 me lope Podlcee'ttoneerenovary 7,57 fs exter digitaren Yoga 1m fies [Emory 50 mers enol 95 {SRyben in ular ory amy exten alle bevy 17 me xen digeorm hen 292,42 roe; te pafrneny 8,95 sb eran 4 flexor hala ben tl leze ptt; #0 Beso iene pee| ‘Muscles ofthe pectoral limb 35 411m, olignassbdnini ext 2 malian sblomiis ns Jim. rcs abdominis; 4m srateveraeabdosnis pectoral Primary 1m, ade ‘ase itil 41, aF ey. prac 28, 28 m, noir 30,39 a sitendinoniss 207 3) ay semeenbranoney 32,92 my. exensoe Cigtrans ioges03 ie ils logos rt i S38, hm. garoenem ar 30 my ener peteatnt 3? me exer, M. extensor digitorum com. (/25); O. Lateral epicondyle of the humerus; of the first and second digits; M. extensor indicis longus (32/14); O.: One head originates from the middle third and the other two from the distal end of the radius and the os carpi radiale; I: First and second phalanges of the second digit; F.: To extend the joint of the second digit; N.: Radial nerve. M, flexor digitorum supf. (32/15); O.: Fascia antebrachii, os carpi ulnare; Lt First and second phalanges of the second digit; F To fold up the wings; Ne: Ulnar and median nerves, First phalanges To extend the finger joints and spread the fingers; N.: Radial Feige M. flexor digitorum prof. (//6); O.: Proxifally on the volar surface of the ulna; Li After ames passing through the tendon of the m. flexor digitorum supf. ics insered on to the recond and wee third phalanges of the second digit; Feo flex the second finger; Nes Median nerves ae M, pronator longus et brevis (/17); O.: Median epicondyle of the humerus; I: Medially cn wae the distal end ofthe radios; F: To pronate the hand; Nu Median nerve, M. supinator; O.: Lateral epicondyle of the humerus; I Dorsal surface of the radius; Fes Supinator of the hands N. Radial nerve. Recondyle of the fad 3rd digits; orans and the surface of the fenaur. I: The es; Ft To flex edial condyle; To extend the : Phalanx T of itexor hallucis iL These are Body cavities 39 Body cavities In birds there is no partition between the cavum pectoris and the cavum abdominis comparable to the diaphragm of mammals. However, aponeurotic membranes, the so-called horizontal and oblique septa, divide the body cavicy into several enclosed compartments which are named according to the organs they accommodate. "The septum horizontale (73/K) is a strong tendinous sheet whichis fused to the ventral surface of the lung and arises from the fused or membranously linked ventral erests, the hypapophyses, of the thoracic vertebrae. It runs in a lateral direction towards the ribs, following the ventral contour of the lung. Te terminates partly by joining the endothoracic fascie and parcly by 2 carved insertion Tine which runs along the ribs, first in a vencral direerion and then ascending towards the pelvis. The eavum pulmonale (73/K), which is a completely enclosed, symmetrical space occupied wholly by the lungs, is limited ventrally by the horizontal septum, laterally by the wall of the thorax and medially by the hypapophyses and the seprum arising from them. ‘A number of openings are present in the horizontal septum which allow connections becween the lang and organs lying outwith this cavity. The main bronchus and the pulmonary vessels enter the lung ventrally, at the point where the septum is fused to che pericardium, Here also are the ostia for the clavicular (/A) and cranial thoracic (Ji) air sacs and somewhat cranial ro these the opening of the cervical air sac (/g). An aperture situated on the lateral border of the septum leads into the caudal thoracic air sac (/l) and near the caudal insertion of the septum on the ribs there is another orifice leading into the abdominal air sac. Four 19 five small muscles knowa as the mm. costopulmonales arise from the endothoracic fascia near the costal insertion cof the horizontal septum. ‘These muscles spread in a fan-like fashion into the seprum and, as individual bundles, atoand the ostia of the air sacs, ‘At the caudal Border of the lung the horizontal septum gives rise to the septum obliquum (73/L). Ac frst ic is attached to che vereebral column whence its insertion line runs in a curve from bebind the last rib, ascending slong the abdominal wall and then ventral to the sternum. Ie is fused cranially with che pericardium. By this means another enclosed space, the cavum subpulmonale, is formed on each side. The cavum subpulmonale surrounds the thoracic air sacs and is bounded dorsally by the horizontal septum, laterally by the thoracic and abdominal wall and ventromedially by the oblique seprum. A chin sheet of muscle, arising from the hypapophyses of the last thoracic vertebrae, spreads into the caudal part of the septum which is situated near the vereebral column. ‘Apart from the bilaterally symmetrical cava pulmonalia, containing the lungs, and the paired cava subpulmonalia, enclosing the thoracic air sacs, there are other compartments in the body cavity of birds for she remaining organs. One of these is limited dorsally by the last cervical and first thoracic vertebra and the cranial part of the horizontal septum, cranially itis limited by the amterior pairs of ribs the coracoid and clavicle and by the skin. In this space are situated the clavicular and pare of the cervical air sacs. The oesophagus, trachea, numerous blood vessels and nerves traverse this cavity. However, by far the largest compartment of the body cavity is che eavum cardioabdominale. It is limited dorsally by the vertebral column and the pelvis, laterally by the two oblique septa and ventrally by the sternum and the muscular abdominal wall, This compartment holds 1.) the heart within its pericardiam, 2.) lying against the sternum, the liver surrounded by the serous hepatoperitoneal sacs (53/1, 1’) and 3.) the proventriculus and spleen. However, the greatest proportion of this part of the body cavity, the visceroperitones! sac (53/3), is occupied by the urogenital organs, the gizzard and the coils of intestine which are flanked by the large abdominal air sacs.“4 ‘Alimentary tract ofthe head has a median and two to three paramedian longitudinal rows of blunt papillae (39/f); in the duds these papillae are confined to the apical region, In the latter species however, there is « median longitudinal swelling. In both the duck and the goose the edges of the palate carry pointed | papillae he mouth cavity snd the rig (0, Floor of the meych and phar: pharyans ef the paoee Sof tbe coe ss spper beak ith Boray lamelte By baie ‘ Pte of he dane, nastics bot PEE nde paso ed bil ‘The following salivary glands (glandd. salivales) are found in che mucous membrane of the palate: 1) The gland. mazillaris is flat and lies against the os incisioum reaching the end of the Palatine clefe. A collecting duct ons almost the whole length of the gland and around it are Prouped the lobules with their mucous terminal cells. The short efferent ducts terminate bilaterally ere apieal region of the palate (37, 39/8). 2.) The mukilobular glandd. palatinae ltt. are situat~ aa cecrel to the lateral ridges in the fowl end their numerous efferent ducts terminate in his Segion (37/}). 3.) Some tubules of the glandd. palatinae medd. are situaced around the palatine see ad others posterior to it. The diseribution of the openings of the numerous efferent ducts Sndicates the extent of the glands (37, 39/R). 4.) The glandd. peerygoidea and gland. tubariae are aaancaod in the raucosa of the roof of the pharynx. The former are more superficial and their efferent ucts open lateral to the infundibular eleft (37/2), whereas the latter are situated in the funnel Sf the auditory tubes, where the ducts also open. 5.) The glandd. mandibulares anteriores liewamedan inthe apical part ofthe floor ofthe mouth, situate in the angle a the pont ofthe lower beak slanduli anguli ovis are situated in the mucosa of the angle of the mouth and they have buc a single efferent duct. Tn all species the mucosa in the region of the auditory ube contains much [ymphoreticular tissue, Lymph follicles, known collectively as the tubat tonsils, are present in the pigeon and are ‘especially abundant in the goose. cs vpee beak: Blower bea mouth cavity drngue phacyagea cai; fei fold; geeophagar arya (atyor FARRER E anal cvs fetal Souay i peatynpll fll» ceulagnour pice of nel vebaey cond nase toma nual pg septum inert) rtdeteplon:» cele with vara medallacerebelarin (5 weds begins pin neds lemeghtlon » meecephlons > pvterys 9 apie sia pharyngeal “The floor of the mouth, which is a ditch-like depression between the rami of the lower beak, : accommodates the tongue (glossa, lingua) (38, 40/¢; 41/d). Ts shape is adapted to that of the lower ‘ray | bbeak and is therefore very variable. In the pigeon the tongue is narrow, whereas in the fowl it ahaa | is broad, lancer shaped and does not extend to the full limie of the lower beak. In Lamellivstres i completely fills the floor of the mouth and is only slightly narrower apically. The body of the tongue contains the of entoglossum. It forms a movable joint with the basihyoid bone which lies fm the base of the tongue and terminates apically in a cartilaginous process. In the avian tongue there is no internal muscle system, analogous to that characteristic structure of the mammalian tongue. Its anterior thied is entirely free of musculature, while muscles of the extralingwal aoe system spread into the remaining parts of the tongue. Its basic structure consists of connective alae tnd adipose tissue and insinuated glands. The free tip of the tongue, which reaches the frenulum Rue Inguae, accounts for the greater part of the total length of the organ and it covers a corres- eee pondingly large part of the sublingual and prefrenular pare of the floor of the mouth. Especially in the apical region, the mucous membrane of the dorsum of the tongue has a heavily keratinised epithelium. In the lower and lateral surfaces and at the base of the tongue, the stratum corneum is of only moderate thickness. In fowls and pigeons the transition between the body and the base ‘of the tongue is marked by a transverse row of upright, backward directed lingual papillae (G8/d). In the duck and goose, in addition to this row of lingual papillae (40/d), there is another row of upright, thorn-like, horny papillae (/d'), situated at the edges of the tongue. They point towards the pharynx and the gaps between them are occupied by sbread-like papillae, This acts asa sieve which effectively supplements the lamellae of the lateral edge of the beak. nate in this the palatine Fferent duces tubariae se their efferent f the funnel mteriores lie Es46 Alimentary tract ofthe ceunk "The tongue contains the glandd. linguales anteriores et posteriores. The former are situated in the lateral surfaces of the tongue and the latter are to be found at the base of the tongue and extend to the larynx (38/e) ‘Organs analogous to the taste papillae of the mammalian tongue are absent from the tongue of, birds: On the other hand, one may find taste duds similar vo those of mammals, singly or in groups, around the openings of the ducts of che glands. They’are seen mainly in the mucosa of the pharyns and on the base of the tongue, and extend as far as the entrance to the oesophagus. Alimentary tract of the trunk (42-53) Foregut ‘The oesophagus (37—53) connects the pharyngeal cavity to the stomach. It is divided into the longer cervical part and the shorter thoracic part. The cervical part is very distensible, specially in aquatic birds. It lies at first dorsal vo the trachea and during its later course turns to the right side of the neck, covered only by skin and accompanied by the w. jugularis dext. Before entering the body cavity between the branches of the furcula, the oesophagus widens to form the crop (ingluvies), a storage organ, which is spherical in the fowl (42—48/b) and the pigeon (46/0) and spindle-shaped in the aquatic species. The thoracic part of the oesophagus (42/a) fs related to the a, carotis dext. et sin, and to the right and left ©. jugularis as it proceeds caudally accompanied by the crachea. It is pushed between the syrinx and the ventral surface of the lung to reach the base of the heart and the dorsal surface of the liver. At the level of the 5rd or 4th intercostal space the oesophagus turns shghtly to the left and ends in the spindle shaped glandular stomach. The thoracie part of the oesophagus is respectively surrounded by the ‘cervical air sacs, the single clavieular air sac and the eranial thoracic air sacs, ‘The muscular coat of the oesophagus is in two or, in some areas, even three layers. Its cutaneous mucous membrane is in longivudinal folds and contains numerous mucous glands, Sithough in the pigeon these are present only in the caudal part. Lymphoreticular tissue is present ‘ind at the distal end of the oesophagus forms into lymph follicles, the so-called oesophageal tonsil Gonsilla oesophagica), whichis prominent in the duck but less well developed in the fowl. "The crop (ingluvies) is capable of storing ingested food for a short time and is an extremely distensible diverticulum of the oesophagus. In the fowl (42—44/b) ic is unilateral and situated ‘on the right but in the pigeon (46/8) ic is bilaterally symmetrical. In the duck and goose, as Siready mentioned, there is no true crop, but simply a spindle-shaped dilatation of the oesophagus. In the fowl it is situated in front of the furcula on the breast musculature; itis covered only by skin and when filled is readily visible and palpable. On its dorsal surface there is the so-called crop channel (fissura oesophagoingluviei) (47/2). Food may go either directly into the stomach bby means of this crop channel or, if there isa larger amount of less easly digestible food, it passes into the dilated crop. In the crop the food is exposed to the action of saliva and probably also regurgitated gastric juices later it is passed into the stomach. Being part of the oesophagus, che trop is lined by cutaneous mucous membrane which, in the fowl), contains mucous glands in the fezion of the crop channel. In the pigeon mucous glands are present in stall numbers only on the veneral surface of the crop. When pigeons are feeding their young, the more highly ascularised crop mucosa of both parents produces the so-called erop milks, This is a crumbly mass Consisting of surface epithelial cells which have undergone fatty change and been desquamaced. Together with other crop contents, this material is used to feed the nestlings. "The stomach (ventriculus, gaster) of the domestic birds consists of two anacomically and functionally distinct parts, the glandular stomach (42/es 45, 48—52/b) and the muscular stomach (aids 44, 45, 48—S2/¢3 53/4). ‘The spindle-shaped glandular stomach or proventriculus (ventriculus glandularis) of the fowl ‘measures some 4em in length and arises directly, without any distinctly demarcated boundary, From the oesophagus. On the other hand, there isa distinct constriction between the proventriculus and the muscular stomach (48—50/2). The proventriculus lies in the lower left quadrant of theesophagus stoinadh ” sitvated gue and congue of, aly or in osa of the ided into sensible, aris dest. videns t0 and the sus (42/a) ayers, Tes glands, is present eal tonsil sxcremely 7 = Alavi B coracid;Csexpla Denes E tid ib Films @ eb Tio sophagus. 1 combs 2 wate, 3 ear abe # etre adtory ments: 5 pper bay 6 lar Beaks 7 extra! aes Vonty by vps ny nme qin nd ey ncn Venn, Me so-called 2 Ee rs etry ih ari ieee sage of ropes odes tayo Hear epee + stomach Ponben hc ony ig Teac som pan “hip aso body cavity and it long ais directed from cranidorslly and medially to caudoventally and agus, the laterally. Ie causes a depression on the dorsal surface of the liver. It is related to the left ab- dds in the dominal air sac and the caudal thoracic air sac of the same side. Together with che spleen, it is only on socommodated in a pocket of the viseeral peritoneal sac. A dorsal peritoneal fold, which carries ‘e highly brandies of the a. coeliaca, envelops the proventriculus and then continues as the ventral ably mass ‘mesentery to the liver. The musculature of the proventriculus has three layers. The mucose is uamated. placed in longitudinal folds and is made up of a single layer of secreting columnar epithelium, Ie contains masses of compound tubular glands which are visible in section with the naked eye sally and Each of these deep propria glands consists of several tubular glands grouped radially around v stomach 1 central collecting duct which terminates in a wide opening on a raised papilla on the mucous rnembrane (50/1). In the neighbourhood of these duet openings there are erypts lined by secreting the fowl epithelium. These erypts are termed the superfieial propria glands. . voundary, In all domestic birds which are primarily vegetarian, the muscular stomach or gizzard (ventri- aes culus muscularis) is constructed, in accordance with its function, as a “masticatory” organ. It is of he Shaped like a biconvex lens and is firm and red in colour. It almost completely fills the left lower48 Alimentary tract ofthe trunk 1 ooh arr # ars; 9» baie se opopharn; beep; « descending and seeding Limb of the doodnemy Fla eeu wean, Et Ppa he Peete Sur ie ny (eh fogs gr ‘quadrant of the body cavity and even extends beyond the midline to the right. The gizzard is related to the dorsal surface of the left lobe of the liver and partly contacts the right lobe. Ceanially and to the right itis related to the spleen; caudally and co the right it contacts various intestinal segments, Tts ventral contour reaches the ventral abdominal wall (53/a), where i is palpable from the exterior. In laying females itis also related to the ovary. The lefe abdominal fir sac lies against the right lateral surface of the gizzard but it can also extend over the dorsal border to the left surface, in as far as this is not joined by connective tissue to the abdominal wall. "The dorsal mesentery, which continues from the proventriculus to the gizzard, carries branches of the a, coeliaca. A serous lamella, which, joins the lig. jaleiforme of the liver, connects the gizzard co the sternum and various segments of the intestine, There is a distinct circular groove at the cranial pole of the gizzard (4850). We differentiate a dorsal (48, 49/5) and a ventral (/6) part of the thick walled body of the gizzard, each with a cranial (/3) and a caudal (/4) thin walled diverticulum. The bright red musculature of the cranial diverticulum (musculus intermedius cran.) continues without any boundary into the dorsal part of the gizzard, while that of the caudal diverticulum (musculus intermedius caud.)Stomich 9 ig. 4. Topography of the neck tna body caviey af she coke fection throagh the nbs; D ‘pectoral uc: Eom loepe eos Fr Pm Toulon, G6 eu senor Shedding nel & aecndng ims of Plone ance Eiake and Bite bes hare 1 aon: 2 a beeocpbalea dex. & continues into the ventral part. Each of these two parts of the body of the gizzard consists of a large mass of smooth muscle which is dark red or bluish-red | in colour, These groups of mus- cles are known as the museuli | laterales (48—50)3", 6°). Their fibres arise from the mother-of- ™ pearl-like endinous aponeuroses (48, 49/7) situated on the lateral sor; 9 a. braioe and medial surfaces. They en aco close the dorsal and ventral Syraid contours of the body of the giz zard on the right side. Nor far from the termination of the “The gizzard is proventriculus is the pylorus # the right lobe. (G9, 50/d) with the origin of the contacts various duodenum, 3/a), where ie is "The mucosa of the gizzard e left abdominal | shows, especially prominently over the dorsal | in the fowl, he zona intermedia abdominal wal (60/2"), whichis a ring of mucous membrane separating it from che proventriculus. Apart from carries branches Iymphoreticular tissue, this zone contains only a few crypts with secreting epithelium. ‘The er, connects the mucous membrane lining the channel-lke interior of che gizzard (50/3', 4’, 8) has a simple columnar epithelium and is arranged in rows of ridgelike folds. The propria contains tabular We differentiate {lands which empty on the surface singly or in groups, according to the species. Despite having ard, each with a 2 completely different function, they have been likened to the pyloric glands of mammals. They sculature of che secrete continuously and the product solidifies on the surface to provide the so-called keratinoid y into the dorsal layer which forms a hard covering on the mucosa. In the region of the tendons, this keratinoid ermedins cand) layer is smooth whereas it forms transverse ridges in the diverticuli (/3', 4”) and longitudinalAlimestary Geral end'= ridges in the remaining ((8). These ac as grinding plates and with the help of che very powerful musculature, they thoroughly trivurate the food, including hard grain. This action of the “masticatory” or gan is effectively assisted by the presence of small stones which grain-eating birds frequently ingest. Small splinters of glass cor even pieces of metal are oceasionally observed in the gizzard and these, like the stones, are gradually ground down. The small and large intestine (4246, 5153) General considerations In birds, as in mammals, one can differentiate the two main divisions of the intestinal canal, namely the small intestine and the large intestine. Morphological characteristics permit further subdivision of the small intestine into the duodenum, jejunum and ilews. The large intestine is ‘very different from chat of the mammal. The presence of two caeca is remarkable and these are(the gander al Bader = ies ofthe pa et Pee ad Pond pe ining regions nding plates » of the very lature, they ie the food, grain, ‘This stieatory” or ssisted by the stones which 8. frequently sters of lass of metal’ are ved. in the se like the vally ground cxtinal canal, ermit further and these are Small and large intestine 51 followed by but a single, shore colon. As well as variations in position and shape, the intestinal segments of the individual species differ in theie relative and absolute length. Ie is very difficult to obtain reliable measurements of intestinal length and those given in the available literature vary considerably, even for the same species. The reasons for this are to be found in individoal, breed and age differences, dietary peculiarities and, finally, the method used in the assessment. The following lise of intestinal lengths can, cherefore, be taken only as average values. The measurements refer to trunk measurements of the birds, taken from the last cervical to the last coceygeal vertebrae. For the fowl this gives values of 1:7—9 =: 165—230em; for the duck 1: 85—I1 =: 155—233 em; for the goose 1: 10—12 = 250—365 cm and for the pigeon 1:5—8 -c71—115 em. The diameters of the small and large intestines are nearly the same although some segments of the gut are characterised by the thickness of their musculature. The duodentm, the ileum and the colon, for instance, have a thicker musculature than the jejunum and the main part of the caeca. The former are, therefore, pale red in colour, while the others are greyish-preen ‘because the intestinal contents show through the wall. ‘The small intestine (intestinum tenue) (42—46, 51—53) From the pylorus of the gizzard arises che duodenum (42/e; 46/e, e's 43/c, c's 44, 48, 51, 52id, d's 53/b, b’). In all domestic birds this forms a characteristic “U*-shaped loop, the straight, parallel limbs’ of which are connected by a strip of mesentery, the ligamentum pancreaticoduo- denale (51, 52/3: 53/5) and embrace the elongated pancreas. The descending, ventral lft limb of the duodenal loop lies against the surface of the right lobe of the liver. Te then runs caudally on the ventral abdominal wall, overlaid by loops of jejunum, turns to the left around che caudal contour of the gizzard and finally makes a sharp bend into the ascending right vencral limb. ‘This also runs on the dorsal surface of the liver, is related to the right testis or the ovary and bends zowards the cranial pole of the right kidney, where it continues as che jejunum. The bile and pancreatic duets enter the ascending limb in a manner characteristic of the species which will be described later. The dorsal mesentery, which has already been mentioned, gives off a double layer of serosa to lie berween the two limbs of the duodenum. Besides this, the ascending limb is connected by a ligament to the gizzard and by another, the lig. hepatoduodenale, which contains the bile ducts, to the porta of the liver. ‘The transition becween the duodenum and the jejunom (42/f; 43/d; 44, 45/es 46/fs 51, 52/e; 53)e) is generally taken to be where the ascending limb of the duodenum bends from left vo right, passing caudal to the a. mesenterica eran. (51, 52/7). The transition from jejunum to ileum, on the other hand, is marked by the liga ileocaccalia, which connect the cacca throughout their entire length t0 the ileum (51, 52/5; 53/0, o'). This arrangement does not apply to the pigeon in arhich the caeca are very short. In the fowl, the duck and the goose the lengih of the ileum is about the same as that of the caeca. A further feature of the ileum, which in this ease also applies to the pigeon, is that its blood supply derives from the 4, ileocecalis; in birds this artery arises, not from the a. mesenterica cran., but from the a. coelica. In birds the jejunum (42/f: 43/d; 44, 45/e; 46/f: 51, 52/e; 53/c) is by far the longest intestinal segment. Its average length for the fowl and the duck is usually said to be 105 em, for the goose 165m and for the pigeon GOcm. Because of its length it occupies most of the right caudal quarter of the body cavity. It is related to the stomach, the spleen, the right lobe of che liver an, in laying females, tothe ovary and loops of the oviduct. "There is 2 marked species difference in the shape of the jejunal loops. In the Fowl (51) there are 10—11 loops of varying size, which form a three-quarter circle around the mesojejunum (51, 52/4; 53/n}, as that part of the mesentery containing the trancus jejunal is termed. In the duck and goose the jejunum is formed into a convolution of 6—8 parallel loops of about equal length, arranged in the long axis of the body. The terminal part of the jejunum and the firse section of the ileum form the supraduadenal loop, so called because of its proximity to the duodenum. Two parts are differentiated in the jejunum of the pigeon (52/e). The first consists of 34 concentric spirals, giving an overall cone effect. Inside this cone there are another 2—3 centrifugal coils. The ascending and descending coils are held together by the mesojejunum, which reaches the apex of the cone and carries branches of the a. mesenterica cran. The second part of the jejunum of the pigeon is the long supraduodenal loop (/e’) which swings to the left and is related to the gizzard.52 Alimentary tract of the tfunk Fig. de Topography of the neck and body cavity of the female pigeon. Let view A ceil; B eres C eal D sens E tid trl probs Om orcobraii “appr bea 2 lower be 3 et ral ausary menue 1 eandiag links of ee duoderas jhe g locas # ans ff of he osophagas Berop dae « eaophagas erp a wah pr The diverticulum caecum vitll, Meckel’s diverticulum, (51/e”) represents the remnant of the yolk duct and of the yolk sac whichis transferred into the body cavity before hatching. This rudimentary struccare will be found as the cupola-like appendage on the convexity of that part of the jejunum which represents the embryonic gut. The mucosa of the diverticulum has deep folds and contains [ymphoreticular tissue, sometimes in the form of follicles. In the dude and goose it persists for along time but in the other species ie disappears eae in life The ileum (42/g; 43/e; 44, 51—53/f) is differentiated from the jejunum by the characteristics already mentioned. Starting neae the cloaca it runs nearly straight in a cranial direction and, inthe fowl, duck and goose, i is flanked on both sides by the eseca, to which itis joined by the ligg. ileocaecalia (51, 52/5; 53/0, 0’). At the level of the gonads ie bends dorsally and to the left and terminates in che large intestine. In the fowl and pigeon the ileum runs along the duodenal loop, in the duck and goose it isin contact with the supraduodenal loop of the jejunum, Icis also related to the right surface ofthe stomachao which eis connected by a serous lamella. Large is 53) estine (intestinum crassum) (42—45, 51 As already mentioned, the large intestine of birds consists of the paired caeca and the colon. ‘The colon is a short length of gut which terminates in a distinct sphincter cloacae where it joins the ampulla-like- dilated first pat ofthe cloaca known as the coprodenm.inane of the ching. This of that part m has deep te dude and rection and, bined by the and to the s along the the jejunum, lamella, d the colon. here it joins Large intétine 33 Figs 48-0, Stomach of the fom tet att rll cae eh lin tion ‘The caeca (42/b; 43, 45/f; 44, 51—53/g) measure 12—25 cm in the fowl, 10—20em in che duck, 22—34em in the goose, bur only 27mm in the pigeon. A circular fold of mucous membrane is present at the openings of exdh of the two cacca where they originace at the site of transition between ileum and colon. They are bound to the ileum by che ligg. ileocaecalia, In the fowl and the aquatic birds, che first pare is short and has a cranial course, then they turn in caudal direction so that their free ends extend to the region of the cloaca. Their relations to neighbouring organs are the same as those mentioned for the ileum. Especially when full, the caeca of the fowl and the aquatic birds show three segments: 1.) the short narrow neck which is very muscular and acts as a sphincter (51/g"), 2.) the long, thin-walled main part (/g"’), whichjena wih the ite Large intestine 58 A 3 ynseotboracic vere: B ams © gums bdecending and ¥ avradag libs ofthe ds id tte at te vers Hepaneer I sighe aed ee dnd (0) fe ete; b colo rc appears greyish-green, because the contents show through and, 3.) the shore but vesicular, bright red apical part (/p"). In the pigeon, where the caeca are only a few mm in length, they appear as bud-like appendages to the colon (32/a, 2"). ‘The colon (45/g: 51—53/h) is a direct continuation of the ileum and is of similar diameter. It is suspended by a short mesentery, which carries the v. coccygomesenterica, a branch of the portal vein, and the a. mesenterica caud. It lies below the vertebral column and ends in the coprodeam. In the fowl the colon measures §—Item, in the duck 7—I2em, in the goose 16—22 em and in the pigeon 3—4 em, The struccure of the tunica serosa, myseularis and mucosa are very similar in both the small and large intestines. The mucous membrane of the entire intestinal canal carties vill, which are particularly long in the colon bue are otherwise of similar structure to the villi of the mammalian Small intestine. The intestine is lined by simple columnar epithelium containing goblet cells which increase considerably in number towards the cloaca. ‘The propria contains tubular intestinal glands, the glands of Lieberkthn. ‘They occur from the pylorus to the coprodeum of the cloaca and they have particularly long tubules in the duodenum, the caecum and the colon. On the other56 Alimentary tract of che trunk hand, the Brunner or submucosal glands, found in the first part of the mammalian small intestine, are absent. The intestinal mucosa is rich in lymphoreticular tissue which is arranged diffusely ot in follicles and even extends into the villi. This lymphoid tissue is particularly abundant in the caeca, In the fowl, duck and goose there is a highly vascularised thickening of the mucosa in che caeca, immediately behind the ned. This is known as the ronsilla caecalis and it contains accumulations of Iymph follicles in crypts. Towards the-cupola-shaped end of the fowl’s cacca, the thickness of the mucosa increases and contains numerous follicles but almost no villi. In the Gack and goose, however, there is a reduction of lymphoreticular tissue in this region. There are xo villi in the very short caeca of the pigeon and the lumen is reduced by mucosal swellings: there is more lymphoreticular than glandular tissue Cloaca (42—46; 51-53 a5 74) In the cloaca, the terminal part of the intestinal tract, three compartments separated by contractile ving folds are recognised. The first of these, the ampulla-like dilated coprodewm (53.alb; 74/k) receives, and temporarily holds, the faeces passed into it from the colon. The second, less extensive, compartment is the wrodeum (/c; /k'). The two ureters (Jb; /m) open into the urodeum in both sexes and\in males the seminal duct (jf; ) and in females, through a slit-like aperture, the Sagitedinal tection of the cloaca of 4 oven fRedrawn ater Clas, 1925) 4 clon; bcopadeus ¢ erodeum; proteus ¢ a f cc sg Sunt Ta hoo 8" oviduct (53 a/i), also open into this segment. The third compartment of the cloaca, the proctodewm (S3ald; 74/k") houses, in aquatic birds, the copulatory organ which is comparable to the mammalian penis. In place of the latter the cockerel has an elevation of the mucous membrane, Finally, the bursa of Fabricius opens on the dorsal wall of the procrodeum. "The somewhat prominent anus (42, 51, 52/i’; 43—45/i; 4614) is closed by the strong im. sphincter ani (53 alf). The bursa Fabricii (53 a/g) develops initially as a solid protruberance on the dorsal wall of the wrodeum. Only when the epithelium-lined lumen develops and forms into a pouch or bursa, does it communicate with the proctodewm. In a fowl of about 8 weeks of age it is round or pear shaped and measures about 1-5-2 cm in length and 0-8 cm in width. With the onset of sexual maturity, it regresses and finally disappears. Its lumen is lined by simple columnar epithelium, ‘Numerous tubules arise from it and between them are lymphoid follicles whose nomber increases during development. Removal of this lymphoreticulay organ leads to 2 reduction of the defense ‘mechanism against infectious diseases. The bursa of Fabricius is sometimes referred to as the “cloacal thymus", for there are certain structural and functional resemblances between these two ‘organs; both contain numerous lymphoid cells and are important in immunological mechanisms.small intestine, anged diffusely abundant in the 2 mucosa in the ind ie contains re fowl’s caeca, no villi. In the gion. There are cosal swellings: sdby contractile (53 albs 74/k) 4s les extensive, rodeum in both apereure, the my a the proctodewm parable to the cous membrane. by the strong dorsal wall of pouch or bursa, 5 round or pear onset of sexual sna epithelium. umber increases 1 of the defense ‘ered to as the ween these two mechanisms. Liver 57 Liver, (Hepar, Jecur) (42—46; 53-58) “The size, weight, consistency and colour of the liver is dependent on the breed, age and the nutritional status of the individual bird. ‘The colour varies from ed-beown through light brown 10 yellow, the latter being seen, for example, in the lipid-rich liver of fattened birds The weights of the livers of different species, both absolute and relative to body weight, are given below: Species absolute weight relative weight Fowl 35— 51 g 1.723 ts of body weight Duck 3:1—#1 %o of body weight Goose 3-6/0 of body weight Pigeon 252-64) of body weight The liver consists of a large right (54—58/b) and a somewhat smaller left lobe (/2). Thee lobes are separated by a deep median incissura caud. ((e) and a shallower incissure cran. (id), with the resule that the two lobes are connected by a bridge of parenchyma. The largest part of the orgen is situated in that part of the body which is enclosed by the ribs, while the remainder, projecting from this space, lies against the sternum. The parietal surface, facies ventralis (55, 58), of the liver is convex and moulded to the contour of the wall of the body cavity. The visceral, dorsal surface (54, 56, 57) is concave. The organs sieuated againse che facies dorsalis, namely che heart, proventriculus, gizzard, the cranial end of the duodenal loop, the spleen and the gall bladder, cause correspondingly named hepatic depressions (54, 55/1—4). On the above parenchymal bridge and continuing onto both lobes of the liver, there is a depression, the fossa transuersa (54, 56, 57ie), which corcesponds to the ports hepatis of mammals. This is where the liver arteries (54/1 56, 37/h, b’) and the branches of the portal vein ($4/b, b’; 56, 57/g, g') enter the organ, and where the bile duces (54/1, m; 56, 57/, k) leave Each liver lobe has sharp lateral and caudal borders and a straight, blant medial border. A small processus intermedius (54/a"", b's 36la’, b’) projeets from the dorsal surface of both main lobes, Figs Stand 55. Liver of the fowl Visceral and parietal aspects Slows meaeven abt and? eke. bepaies ea 1 inpreio ealsy 2 imprenia aden; 3 peeing; # impress/ 58 Liver Pints, Liver of che goose. Visceral aspect t eft lobe with pac insermedian (2) B ihe Lbe wit roc inermeine Ge fou fonaey b sight and Ba a he Mens sp oblieraned vs wmbibeais i baer, caudal to the transverse fossa, The 9. cava caud. (54, 55/g; 56—58/f) tunnels through | the cranial part of the right lobe of the liver, forming a caval canal, | "The liver, covered by a thin fibrosa and | the serosa, is accomodated in the hepatope- | ritoneal sacs (cava peritonaei_hepatis) | Galt, 7), Coresponng to the division of {his organs we differentiate large ht and left vental hepavopeitoneal sae for tach ofthe main lobes and smaller doreal tec for cach of the invermediae process \ ‘These hepatoperitoneal sacs reach the hori- | omal sopeumeanilly adits Stree, the vevural surface of the’ lung nd te gonads All the aie sacx apart from the cervical, are in contact for a variable ement with the hepatopercones sacs, Two Hgumens, which corespond tothe liga, tianguaria of mammals, un beween the caow Der tonaei doysale and ventrale and attach the right and the left lobes laterally to the wall of the | oy cavity. A double serovslamll,aing on the dorsal wal of the abdomen ata conination OF the mesenterinm dorsal, is inserted in the fossa tansversa of the liver and it continues, | lee ea i ae epee ep mean $Simown asthe lig. flesforme (33), / ‘The livers of te diferent Species of domestic birds have certain charaternc diferenes in shape, some of whish are worth recording here. For examples in the fowl there is nor much difference in size between the ellipsoid left and the nearly heart-shaped right lobes: The incissura intvalobularis (54, 55/f) which divides the left lobe into 2 lobus sinister lat. et med. is very - 4 tele lbe; ihe Tobe ¢ foe wanereay 4 Jnciarn crams # icra cas ff 7 cava cud g ge and ethos and es hpacns | duces beptontercs ses Feces Heotcncas detSta pe + through ve of the vrosa and aepatope- hepats) ivision of ge right Voss for ler dorsal processes. the bori- thos, in- the’ lung tes, apart sacs. Two ‘eum perix all of the mesentery, not much d. is very sed of Be Gall Bladder 59 characteristic. The pear-shaped gall bladder is embedded in the caudal half of che right lobe and ies apex extends to the caudal border of that lobe. In the dick, on the other hand, the right lobe of the liver is tongue-shaped and remarkably long whereas the left lobe is more bean-shaped. In he goose the liver as a wholé appears broader and again the right lobe is longer than the left. In both the duck and the goose the gall bladder is situated in about the middle part of the dorsal sucface of the right lobes it is elongated and tube-like aiid does not extend to the caudal border Of the lobe. In the pigeon there is neither an intermediate process nor a gall bladder. ‘The liver of birds ws constructed as a tubular gland, che parenchyma of which consists of tubes of cells connected together in a net-like fashion. The axial lumina of these cubules are the bile Capillaries, The wall of the tubules is made up of liver cells between which are situated the ftercellular bile canaliculi. In eross section these tubules consist of between two and six cells. ‘Closely interwoven with the tubules are the capillaries, or sinusoids, of the afferent portal system, associated with which are the Kupffer cells. The capillaries collect in veins, situated in the centre of each complex of tubular glands, which is analogous with a lobule, These central ‘ring continue into the sublobular veins and finally, through the ow. hepatica, from the ©. caus aud. "The bile canaliculi feed the intrahepatic bile ducts which, in vurn, unite to form the extra hepatic bile ducts. The simple prismatic epithelium of the latter is devoid of goblet cells. The large bile ducts and the gall bladder contain neither goblet cells nor glands. During embryonic life the liver is an important site of blood formation uncil this function is taken over by the bone marrow. ‘The liver is concerned in the break-down of haemoglobin and its conversion to bile pigment, It thus carzies a considerable store of iron. As a producer of bile, the liver functions asa secretory gland but by supplying the blood with substances synthesised ‘and stored in the liver cell its function might be compared to hat of an endocrine gland. "The imporsance of the liver to the entire metabolism of the body can be recognised by the fact that ir receives through the portal vein the bulk of the substances absorbed by the gut. But since the substances passed to the liver during digestion are in such a quantitative and qualitative state that they eannot be utilised directly by the body, they have to be transformed and stored in the liver, They ean then be released from the liver into the blood as required and transported to the site of utilisation “among the functions of the liver as the central metabolic organ, one might mention trans formation of blood glucose into glycogen and, after temporary storage, the release of dextrose Sato the blood, Excess cazbohydeates ean be converted to fats and stored in the liver cells. The liver also plays a decisive role in protein metabolism, converting the amino acids supplied t0 Heinto the protein characteristic of the species. Alternatively, it can urilise amino acids for conversion t0 glucose. By conversing noxious substances taken up by the gut, or even produced in the body, into harmless products which can then be eliminaved, the liver plays an important role in detoxication aad defense, This function includes the conversion of the protein break-down products into sine acid, Farthermore the liver, as an important component of the body's reticuloendothelial ‘Stem, produces antibodies for defense against infections. “The gall bladder (vesicafellea) (54/R; 56i!) lies in the fosca vesiae felleae on the dorsal surface of the Fight lobe of the liver. It is partly fused to the liver parenchyma and partly covered by peritoneum, In the fowl itis pear-shaped and its caudally directed apex extends to the caudal Perder of the right lobe. In aquatic birds it is more tubular in shape and does nor extend as far te the caudal border. ‘The following extrahepatic bile ducts can be identified: 1.) The ductus bepatocystci which empty into the gall bladder, 2.) The ductus hepatoentericus (S1/1" 4/43 5¢i) catries the bile directly into the duodenum and 3.) The ductus eysticoentericus (51/1"”"s 3a/mn; 36/k) runs between the gall bladder and che duodenum. In fowls and water birds the two jase named open into the ascending limb, of the duodenum. Tn the pigeon, which has no gall bladder, there are only two dactus bepatoenterici (54/1,1'; 57/i, /k) and the sight duct empties nwo the ascending limb and the left, and larger, duct into the descending limb of the duodenum,60 Pancreas Pancreas (43, 531; 48/0; 46.5 51, $2/n) ‘The pinkish, ribbon-like pancreas of birds is situated between the two limbs of the duodenum and enclosed within the lig. pancreaticodwodenale (51—52/3; 53s), a double serous lamella which connects the two intestinal segments. We can differentiate a dorsal and a ventral pancreatic lobe (lobus dors. et ventr.) (45im, ms 51, 52/n, n’) as well as a very narrow strip of pancreas running towards the spleen, which is particularly rich in isles of Langerhans, and is known as the splenic lobe (51, 52/n”). In some cases this thin splenic lobe, which is embedded in the adipose tissue, can barely be seen with the unaided eye but eis always demonstrable microscopically. In the fowl and pigeon the pancreas almost completely fills the gap between the limbs of the duodenum, whereas in the duck and goose the duodenal loop extends well beyond the end of the pancreas. In the fowl, and sometimes also in the goose, the dorsal and ventral lobes of the pancreas are connected by a parenchymal bridge of varying width. In the duck and pigeon and sometimes the goose, the two lobes remain independent. In the duck, goose and pigeon the splenic lobe is connected only with the dorsal lobe while in the fowl ie joins both the main lobes. ‘Asa rule there are three efferent pancreatic ducts in the fowl and the pigeon, two of which arise from the ventral (51, 52/2) and one from dorsal (/2') lobe. Usually there are only two ducts in the duck and goose although three sometimes occur. In all species these ducts empty into the ascending limb of the duodenum. ‘The pancreas is made up of two functionally quite distinct components. The bulk of the organ consists of secretory glandular tissue whose product, the pancreatic juice, contains che following enzymes or proenzymes: 1.) the proteolytic proteases trypsin and chymotrypsin; 2.) the starch digesting enzyme amylase and 3.) the lipolytic lipase steapsin "The secretory part of the pancreas has the structure of a compound tubular gland whose secreting terminal cells are bound into lobules by loose connective tissue. The thin, serous pancreatic secretion is produced by the glandular cells from zymogen granules and caeried into the duodenum by two or three duces. “The second component is the endocrine islet tissue, which is made up of microscopic groups of cells embedded in the glandular parenchyma and known as the pancreatic islets or the islets of Langerhans. They produce two hormones, insulin and glucagon, which together with other hormones have a decisive influence on carbohydrate metabolism. ‘The pancreatic islets, rich in blood capillaries, are made up of A-, B- and D-cells which are morphologically and tinctorially different. The production of glucagon is attributed to the Accells and insulin is produced by the B-cells but the function of the D-cels is still uncerain, A peculiarity of avian pancreatic islets is that some consist almost exclusively of A-cell, which are then referred co as the dark islets, whereas others are made up predominantly of B-cells and are known as light islets. Finally ie should be recalled that the splenic lobe contains very many isles Spleen (lien, splen) (51, 52/m) ‘The spleen is a small organ of brownish-red colour. In che fowl ie is round or egg-shaped, in the aquatic birds more triangular with a flattened dorsal and a convex veneral surface, and in the pigeon itis oval. The spleen of the fowl and duck weighs 1-5—45 g, that of the goose 4—8 g and that of the pigeon 02—0+4 g Ie lies against the dorsal surface of the right lobe of the livet in a space formed dorsally by the gonad, ventrally by the liver and laverally by the gizzard. From the mesenterium dorsale it receives a lamella of serosa, which continues onto the proventti= culus and contains branches ofthe 2. coeliaca Structurally and functionally the spleen belongs to the blood-forming organs and at che same time it is also an important component of the feticuloendothelial system which serves as a defense against noxious substances. In the embryo it produces both red and white bood cells and it retains lymphopoietic ability throughout life.duodenum nella which seatic lobe sas running: the splenic pose tissue, imbs of the end of the shes of the pigeon and Pigeon the ‘main lobes. which arise vvo ducts in ty into the £ the organ « following the starch and whose hin, serous sartied into «groups of he islets of with other 5 which are nted to the calls, which Beeells and very many ssthaped, in face, and in ose 4—8 g of the liver he gizzard, > proventri- at che same bood cells Spleci, 6t ‘The spleen also removes erythrocytes which are no longer viable from the circulation. Some components of the blood pigment are passed to the liver for further processing, while iron is deposited in the spleen in the form of ferritin. "The phagocytic reticulum-cells, which form the basic structure of the splenic parenchyma, are concerned mainly with the above functions and with defense against infection. ‘The endothelial cells of the splenic sinus have a similar function. Bork these cell types are also concerned with antibody formation. Furthermore, che reticulum cells are the stem cells of the lymphocytes. The connective tissue capsule of the spleen is covered by peritoneum and forms connective tissue trabeculae which enter the parenchyma, carrying with them the larger blood vessels. The framework of the parenchyma consists of reticular connective tissue which contain the splenic sinuses, fed by arterial capillaries and drained by veins. Because of its high blood content and its soft consistency, the main part of the parenchyma is known as the red splenic pulp. The arteries of the pulp are surrounded by sheaths which consist of a dense network of reticulum cells, the meshes of which contain many lymphocytes. ‘The lymph sheaths of the arteries are sometimes enlarged to form small lymph nodules which show distinct reaction centres. The number of these so-called splenic corpuscles varies according to the reaction seats of che organ. They are collectively spoken of as the white splenic pulp,choanal cleft floor of the the cartilago and epigloutis woalar ventral us or ossified sing the semi noid cartilage cartilagenous age, is covered of keratinized fous membrane sified ciliated = width of the ns lat. acts a8 border of the medial t0 the the arytenoid of the larynx, of muscle and, covered only. ‘wl and pigeon of the fureula, rminal part of or larynx cad. of the trachea the neck and seen the rings hat the trachea "The trachea is achealis (441F), Syrinx, tung 65 arises from the sternum and the ventral angle of the furcula and the other, known as the im. sternotrachealis (44/G; 451C), from the cranial process of the sternum. In the drake, about 6 tracheal rings of the bifurcatio tracbeae fuse to form a thin-walled bony vesicle which projects mainly to the left, Ie is known as the Bulla tympaniformis (62, 63) and is an organ of resonance, which increases the voice volume. "The syrinx or larynx caud. (64) is situated at ahoue the level of the 2nd or 3rd cervical vertebra and is enclosed by the clavicular air sac. As already mentioned, it is made up of the terminal part of the trachea and che first part of the ewo main bronchi. The part of this syrinx tradheobrondhalis known a8 the tympanum (Ja) is constricted laterally in the fowl and slightly dilated in the duck and goose. The wall of the tympanum is formed by the last, fused tracheal ings. In the tympanum between the two openings leading into the main bronchi there is a sagitally placed, comb-like proteuding beam of cartilage, the pessulus (/b). Its free border may carry a sickle-shaped fold, From both sides of the pessulus arises an elastic, mucosa-covered membrane, the membrana tympaniformis int. (/e). This membrane forms the medial wall of the, {pselateral main bronchus. The membrana tympaniformis ext. (fd) is stretched between the outer ‘wall of the tympanum and the first bronchial ring. In analogy with the vocal chords of mammals, these membranes are activated by musculature which is highly differentiated in singing birds. In domestic birds, however, there are only the two muscles previously mentioned, the m. ypsilo~ trachealis and the m. sternotrachealis. Fi.4l, Sysinx (grams) satisfactory birds whose reproductive activity ation in birds follows a strict rhythm adjusted to to the organ climatic conditions. Thus, for example, the volume of the sparrow’s testis varies between the size of a pin head and that of a cherry; this represents 1 volume increase of 300 times. The proliferative and regressive changes affect the sperm-forming tubules of the tests. ‘The testis is yellowish to greyish- white during the resting stage and almost pure white during the stage of activity. Te is enclosed in the fi- brows tunica albugines and external to this it is covered by peritoneum. Delicate tracts of connective tissue portal system uchors believe metabolism te sive of their veloped their radiate from the tunica into the or- ‘only they ex- gan but true septa are not formed. related t0 the Pigment cells are not infrequently rxact with the found in fairly large numbers in the and the ab- connective tissue capsule and the sal abdominal intersisium of the testes in both the ves. The testes ' cockerel and turkey stag. The very twisted seminiferous cubules (tubuli seminiferi contorti) are cies, they are embedded in the vascular interstitial connective tissue which also contains the hormone-forming s. During the ‘Leydig cells, ‘These seminiferous vubules, which measure 150—200 um in diamever, are enclosed nin diameter by a thin tunica propria which consists of collagen fibres. Internal to this is the basal membrane mm long and ‘on which lies the germinal epithelium made yp of two functionally different cell types. These are is has a lengch the supporting Sertoli cells and the germinal cells. The Sertoli cells have an oval nucleus, poor in and 9—21 mm. chromatin, Their broad base lies against the basal membrane and the slim part of their eytoplasm 34mm thids projects into the lumen of the tubules, spreading out in finger-like processes, Between these Ser- fe can measure toli cell all the different developmental stages of the germ cells may be seen extending’ up vo the74 Sex organs lumen. Spermiogenesis is the process of development of these germ cells into mature spermatozoa. ‘The seminiferous vabules collect into the rete testis. Arising from these are the ductuli efferentes and, subsequently, the ductus epididymidis and together these two form the small epididymis situated on the dorsomedial aspect of the testis. Spermiogenesis: Immediately on the basal membrane of the seminiferous tubule are situated the spermatogonia which arose from the primordial germ cell and have a spherical, chromatin-rich nucleus. These divide equatorially giving rise to two unequal cells. One of these cells remains a spermatogonitum while the other develops into a primary spermatocyte. ‘This bivalent multi- plication ensures the persistence of the spermatogonia and therefore the continuing formation of further primary spermatocytes. The primary spermatocytes grow into large cells wich loosely structured auclei; there is pairing or conjugation of homologous chromosomes allowing the ex- change of genes. In the first maturation division the primary spermatocytes divide by reduction division or meiosis, without preceding longitudinal cleavage of their chromosomes, into two daughter cells the secondary spermatocytes, which have haploid chromosome complement. Two haploid spermatids, the precursors of the spermatozoa, result from the second maturation division, which is a normal mitotic division, of the secondary spermatocytes. Thus from one primary spermatocyte four haploid spermatids have developed. These are strikingly small cells, the apical ends of which are in close contact with the Sertoli cells, thus facilitating the exchange of nutrients. ‘The spermatids ‘now undergo the process known as spermiogenesis which means the transformation of spermatids into spermatozoa. After completion of this transformation process, the spermatozoa consist of a head, neck, midpiece and rail, ‘Alchough from the point of view of volume the sperms are among the smallest cells of the body, they are of considerable length. For example, the sperms of the fowl measure 90—100 ym, while those of the pigeon are 180 jum in length. The longest part is the rail. ‘The sperms, which at this First stage are still immobile, are carzied by the stream of secretion from the seminiferous tubules to the reve testes, thence to the ductuli efferentes and finally through the ductus epididymidis into the ductus deferens, where they complete their maturation. The ductus deferens arises, without any clear line of differentiation from the epididymis (45g; 74/f). Te accompanies the ipselateral ureter beneath which it crosses from medial to lateral at the level of the caudal lobe of the kidney, to terminate in a small papilla in the urodeum (Sali; 74/l), the middle compartment of the cloaca. Daring the mating period the ductus deferens increases considerably both in length and thickness and becomes folded into numerous meandering convolutions. At that time ie acts as a sperm reservoir and is recognisable by its white colour, The epithelium of the epididymal canals produce a secretion, but such function has not been demonstrated in the epithelium of the ductus deferens. The male bird has no accessory sex alands: ‘The semen or ejaculate consists of the spermatozoa and the epididymal secretion. In the fowl the ejaculate has a volume of between 0-8 and 1-0 ml and the number of sperms lies between 550,000 and 6,000,000 per emm. The concentration of the semen decreases with the number of copulations. A cockerel can copulate, or teead, up t0 50 times per day. Once the sperms have been introduced into the female genital tract, they remain fertile for 10—21 days in the fowl and 8-10 days in the duck and goose. Male copulatory organ (penis) ‘The copulatory organ, or penis, is rudimentary in the cockerel and absent in the pigeon. In both species the everted proctodeum accomplishes the transfer of semen into the cloaca of the Female which is also everted during mating. In contrast, the gander and drake (45/i) both have a copulatory organ which in ehe erect stave, measures about 6—8 cm in the drake and 7—9.em in the sander. The penis arises at the transition between urodeum and proctodeum on the ventral surface Of the cloaca. Ie consists of a short, connective tissue body and a free tube, reaching to the tip of the penis. The organ is covered with mucous membrane and it forms a seminal groove in spiral coils. At the edges of the groove the mucous membrane is raised by the presence of erectile tissue During copulation this groove is closed forming a tube which transfers the semen into the cloaca or the terminal part of the oviduct (vagina) of the female. The penis is protruded from andefferentes epididymis ivuated the vmatin-rich ent multi- mation of ith loosely rng the ex- Jivision of er cells the permatids, sa normal ocyte four which are spermatids spermatids consist of f the body, dyum, while hich at this ‘ous tubules ymidis into epididymis Ito lateral fe urodeum the ductus vy ite white fon has not cestory sex fn the fowl es between number of shave been = fowl and pigeon. In aca of the doth have a 9 cm in the tral surface o the tip of ve in spiral zetile tissue > the cloaca i from and Female sex organs 6 retracted into the male’s cloaca by muscular action. Erection of the organ is achieved by the influx of lymph into the interior of the blind cube. This fluid is supplied by a pair of Lymph- ‘producing cavities situated at the base of the penis, which, in their turn, are in functional relation to the so-called vascular bodies from which the lymph permeates. In newly-hatched male chicks a rudimentary copulatory organ is present on the ventral wall of the cloaca. It consists of 2 basal part and a tight, shiliy upper pare which is pointed or convex, With appropriate dexterity, this small raised structure can be everted from the cloaca and the sex of the day-old chick chus determined with a high degree of reliability. Female sex organs (organa genitalia feminina) (42, 43, 46, 51—53, 7581) Ovary Although there are two gonads during developmental stages, in the majority of birds only the Jefe ovary (42/5; 43/m; 73ley 76g: 77/73 78/a) reaches fll development. Inthe fowl development of the right ovary has already fallen behind the left by the 7th day of incubation and within the firse days after hatching i has all disappeared except for 2 small remnant. ‘Shortly before hatching the ovary of the chick measures 56mm in length, {-5—2mm in width and 0709 mm in depth. Tris yellowish white and the shape of an elongated triangle. The surface, covered by germinal epithelium, is smooth, The ovary lies flat against the anterior lobe of the left kidney, reaching the adrenal gland with its anterior border and the aorta medially. On the lefties related to the ipselaeral abdominal air sac and ventrally its related tothe stomach “The prepuberal phase of development of the ovary stares after hatching and continues until the onset of the first laying period. During this period the size increases and the surface develops characteristic contours although the gonad is still non-functional. Because available space is restricted and because the ovary is so closely attached to its blood supply, transverse ridges, Separated by grooves of varying depth, develop during its further differentiation and growth. In 2 twormonths old pullet the ovatian surface is finely granular buc by che beginning of the tied ‘month individual ripening follicles ean be clearly differemiated. From then onwards theiz growth is very rapid, so that by the end of the third month they already measure O-5—1'5 em in diameter. By the time the hens come into lay at an age of about 5 months, depending on the breed, the follicles gradually grow to 5:5—4 cm in diameter. The smaller follicles are greyish-yellow, the larger ones whive and those above 5 mam in diameter are yellow in colour. The ovary of mature hens (75/¢} 77/75 7812) bas, besides a large number of follicles with a diameter of 310mm, a number of dark yellow follicles measuring bevween 5 and 10mm and even up to 40mm ia diameter In all species the lacter, large follicles are distincly stalked and the enveloping membrane ‘ies numerous blood vessels except on a sickle-shaped strip, the stigma (75/1), which lies on the eurface opposite the stalk. The numerous, maturing follicles which exhibit a wide range of sizes, result in a considerable increase in the size of the ovary and a change in is form. In day-old Leghorn chicks the ovary weighs 0-03 g bue ie reaches a weight of 246 g in 4-month-old pullets and 38 in laying hens. In appearance it could be compared ro a bunch of grapes, with the stalked, large Follicies hanging into the body cavity. The ovary has now reached the stage at which gg production can commence. At ovulation che follicular membrane ruptures atthe avascular sit, the sigma, and the oowm or sphere of yolk (78/1; 771r'; 78) is liberated into the infundibulum ‘whichis the fnne-shaped, fest part ofthe oviduct. The follicular membrane, which is now empty, is lefe behind as the so-called ealfx (75/25 78/2), In the fowl this represses completely within 10—14 days or, as recent investigations have shown, even within 7 days. Tn wild bieds and the less highly-bred domestic fowls, as well as all other domestic birds, snaturation of the follicles and ovulation cease completely as egg laying is interrupted by the ‘onset of Broodiness and the moult, In modern laying breeds in which the incubation drive has been eliminated by genetie selection, ege production is interrupted only by the moult. This biologcally- determined pause in the laying cycle is initiated by regressive changes in the ovary which are macroscopically discernible. The first noticeable change is retarded growth of the maturing follicles, This is followed by resorption of both the large and the smaller follicles so thatthe ovary resumesecht Oogenesis 7 the appearance and size of the prepuberal state (76/g). In the fowl such a laying pause lasts 2-5 months during the moult and it recurs with great regularity in older bieds, The time of onset land the duration of the moult, and therefore of the laying pause, can be influenced both by selection and by diet and husbandry. This applies to the fowl and to other domestic birds, If there is only a short pause in the ovulatory cycle, then regression and the production of new follicles go on together, so that, after a relatively"short time, new ova are discharged from the ovary. ‘The surface of the ovary in birds, as in mammals, is covered by a single layer of germinal epithelium which is partly cuboidal and partly columnar and is situated on a thin layer of ‘connective tissue. Internal to this is the stroma, and these layers comprise the cortical zone (zona parendhymatosa). The cortex surrounds the medullary zone (zona vasculosa). The follicles con- tained in the cortical zone consist of the oozyte and the single layer of the follicular epithelium surrounding. it, which is columnar at first but becomes more flattened later. The follicle wall also contains the fibrous theca folliculi which is very chin at first but later increases in thickness and becomes double layered (theca externa and theca interna). It contains numerous blood vessels. The medullary zone is made up of loose connective tissue containing numerous blood vessels and lymph spaces and a few muscle fibres which extend into the cortical zone. Groups of interstitial cells, to which are attributed endocrine functions, are mainly situated in the cortex but also occur in the medullary zone. Formation and maturation of the ovum, oogenesis During early embryonic development the so-called primordial germ cells migrate into the gonadal anlage and differentiate, in the female, into primitive egg cells known as oogonia, These smuluiply together with cells derived from the germinal epithelium and they form the germinal cords of the cortex. Subsequently they are divided by vascular connective tisue strands ino cellular complexes which then give rise to the follicles desribed above. Tt is not uncommon t0 find follicles which contain two or three ova. In birds four to eight days of age this process of follicle formation is nearly completed and in the fowl the follicle size is usually between 10—25 jum but may be up to 50 um. This completes the multiplication period of the gg cel With the onset of sexual maturity, the oogonia grow into primary oocytes. Duting the first phase of this growth period the nucleus of the ovum develops to its ultimate size, while the éytoplasm increases only slightly. During the second phase there is a striking increase in the concentrically layered sphere of yolk. In the macure state che eg eel, surrounded by a transparent membrane, shows the following structure: at its animal pole there is the peripherally situated germinal disc (7918) consisting of the flawened call nucleus (germinal vesile) surrounded by a small amount of cytoplasm, The germinal dis lies upon the latebra (Je) whichis made up of white yolk. The centrally situated latebra, which has a pegrlike extension to the surface, is sucrounded by alternating layers of pale and dark yellow yolk, which make up the sphere of yolk (!d,e). Fxg production is governed by the rhythm of follicle maturation. A hen in intensive lay will ovulate daily and a drop in production isthe result of retarded follicle growth during the second phase. With the onset of sexual maturity, the oogonia grow and their yolk content increases to form the primary oocytes. This growth period is followed by the maturation period in the course of which the ripe ovulum develops and is ready to be fertilised. This is achieved by ewo maturation divisions. After dromosome pairing and exchange, the cell nucleus, the so-called germinal vesicle, moves towards one pole of the primary oocyte and one half ofits chromosome complement, together with a small amount of cytoplasm, is pindied off as the polar body ot polecyte. The cell has now become a secondary oocyte. At the moment of fertilisation when the sperm enters the ovum, the secondary oocyte goes through another similar maturation division, eiving rise to a second polar body. At the same time the fist polar body also divides and the result of these meioses is three haploid polar bodies and che mavire ovum, which is also haploid ‘The three polar bodies de; they simply dissolve in the mass of yolk. ‘The fundamental differences in che sex or XY-chromosome constellation of mammals and bieds should be noted. In mammals the cromosome formula for the male is 2n+xy, and for the female 2n + xx. Therefore, in respect of its sex chromosomes, the mammalian male is heterozygote andOviduer 79 the female homozygote. The sperms are either n + y (androsperms) or n + x (gynaecosperms); the ovemis invariably n + x, In birds the situation is reversed in chat the female has a chromosome complement of 2n ++ xy and the male 2n + xx, so that the female is heterozygote and the male homozygote. The chromo- some formula for the mature egg is therefore either n + x (male ovum) or n+ y (female ovum) while all the sperms have a chromosome complement of n + x ‘The sex of the individual is thus determined by the sperm in mammals and by the ovum in bieds "The number of follicles intially present in che fully developed ovary far exceeds the number of eggs produced during the several years of laying activity. The face that in old birds the ovary is completely devoid of follicles is due to a process described as follicular atresia. This is degeneration of follicles of all sizes and in all functional stages of the ovary. The process is initiated by multiplication of the blood vessels of the theca folliculi and atrophy of the follicular «epithelium with simultaneous absorption of the content of the follicles. Nests of epithelioid cells remain in the ovary as the remnants of such atrophic follicles they are enclosed by fibrous tissue ‘The larger follicles sometimes empty their contents into the surrounding connective tissue, where they are absorbed. ‘Numerous investigations have been concerned with the question as to whether the follicular capsules, the calyces, proliferate after ovulation and form structures which produce progesterone. If this occurs they would be comparable to the corpus luteum of mammals. Some authors believe this to be s0, bur more recent investigations have shown chat there is no corpus buteum in birds. It has been demonstrated that degenerative processes take place in the follicular capsule after ovulation leading to complete dissolution within a few days Reference should finally be made to changes in the gonads of fowls which result in some cases of sex reversal, It has been stated above that the right ovary lags behind che left in development. ‘The rudimentary remains of the right gonadal anlage consists of medullary tissue, further development of which is suppressed under normal conditions by the female sex hormone produced in the cortical zone of the left ovary. If, in old hens, nacural exhaustion of the corsical zone of the previously intact left ovary leads to loss of female sex hormone production, then the rudiment of the right gonadal anlage may develop into a testis. A similar resule may occur if here is pathological destruction of the cortex of the left ovary. This sex reversal not only induces the formation of male plumage but both in appearance and behaviour such individuals are like true cockerels; they are even able to fertilise other females. Oviduct Because the anlage of the female avian gonads is paired, there are two Millerian ducts during embryonic development. However, only the left grows into the oviduct while, as a general rule the right entirely regresses. In fowls one may occasionally find non-functional rudiments of the right oviduct but these lack patent communication with the cloaca. ‘Synchronous with the changes which occur in the ovary under the conteol of the gonadotrophic hormones of the pituitary, there are fundamental cyclic changes and functional modification in the oviduct. These are initiated and subsequently controlled by the ovarian hormones oestrogen land progesterone. The term “oviduct” gives only an incomplete indication of the pare played by this complex organ in the manufacture of the avian egg. Essentially it provides the fertilised ovum with the additional nutrients and protective membranes necessary to safeguard embryonic development outside the maternal environment. Tn a day-old chick the oviduct is present as a tiny tube just visible to the naked eye. This gradually increases in size until puberty and in the laying hen it is a 60—70em long, gut-like trgan. At this stage the oviduct is more than double the length of the bird's trunk and i reaches from the ovary to the cloaca in a massive coil. It fills the left upper quadrant of the body cavity completely and also extends into the lower quadrant of the same side (42/3 53/m; 73; 77). Its inicial funnel-shaped part, the infundibulum (78/d; 78/6), is pushed between the maturing fol- licles where itis fixed by a stout ligament extending from the second last rib on the left side (78/4). The remainder of the oviduct is held in place by a shore peritoneal fold, the mesoviductumSex organs Fig 7h Ovacy with che Siete part of she oviduct of © Tnying hens Venel view cfeadibalua, co. oF sing ao ep cl fm aio of eying ages 3 afd is ‘iia of dhe infendielams 9 ena stpeoey fpakehtar te vic: € dn gnnent of the ode (75/7; 7815, 16), which contains muscle fibres and numerous blood vessels, and is inserted onto the left kidney and the dorsal abdominal wall. ‘In broody hens the total length of the oviduer is re- duced t0 about 30m, while during the full moult itis only about 18 em long and is about as whick as 2 straw Te regains its larger size during the next cycle (76). ‘The pale pink wall of che oviduct consists of three layers. Externally it is covered by peritoneum, then fol- ows a double layer of smooch muscle and finally the lumen is lined by a very vascular mucous membrane. "This mucosa has such tall folds that they may appear 10 fill the lumen. Numerous studies have been carried out on. the morphology and function of the mucosa of the oviduct land the formation and structure of the egg membranes The fully developed oviduct can be divided into five regions: 1.) the fonnel of the oviduct, the infundibulum (id; 78/b), 2) the oviduct in the strict sense, che magnum (75/ei 77/5: 78/4), 3.) the macro [ncof the oviduce, che thus (75; 77/0) 4) the sll gland or wteras (75/g5 77/H) and 5.) the Ja (75183 7/9). In laying hens the average length of the infundibulum is 7+ em, the magnum Sam, the isthmus &-7 em, the uterus 8:3 em and the vagina 7 em. The hinwalled, funnel-shaped infundibulum has an opening about &.cm in diameter and it is connected to the magaum by a short tube (78/c). In the first part of the infundibulum the mucosa foams low, closely packed folds which take a slightly spiral course. Lower down these folds increase tn height, The mucosa of the magnum has tall, coarse folds also arranged in a spiral. There is = io poh anitional area between the magoum and the slim, eube-like isthynus where the folds are ery Tow, but they cepain their original height distally, However, in this distal part of the ithrios Wrholde resemble flat leaves, the lateral surface of which carry small secondary folds. The profile OF he ride sterns differs only in as much as the folds, which are of constant height, appear much saarer in cross section. In this region there are also circular ridges formed by the musculature ratio all, The mucosa of the vagina presents a similar picture. This terminal segment of the Sriduer opens into the middle part of the cloaca, the wrodeum (53./es 77/w), via a slit-like opening. "The snacroscopic appearance of the oviduct suggests that the individual regions of this organ perform different fonctions. But a stody of the histological morphology of the mucosa in the Ae Hoont regions is necessary to obtain an understanding of the processes which occur in the oviduct Seen the fits entrance of the ovum until the end product, the completed egg, is ready to be Inid ton Nonnection, particular attention must be paid to the microscopic structure of the epithelial surface of the mucous membrane and the glands of the mucosa. TA Tint the mocous membrane of the infundibulum is covered by a single layer of fla epithelium br ahis geadually becomes columnar. In the tubular part of the infundibalum, the epithelium has Enanged invo a multilayered, serering type whichis partly ciliated. ‘There are no glands in the sarees membrane of the initial pare but in the more distal region it contains first vesicular, and Tee branching, tubular glands which have a proteinaceous secretion. In the magnum the multi- 1noced, columnar epithelism changes to a single layer consisting first of cuboidal, and then ofoviduct of # » fibres and ‘onto the lefe oviduct is re al moult ie is asa straw. (76). sists of three cum, then fol- id finally the as membrane. ‘ay appear to out onthe of the oviduct ig membranes. ded into five infundibulum ) the narrow i) and 5.) the ‘the magnum acter and it is tm the mucosa folds increase sal, There is a fe the folds are of the isthmus 4s, The profile > appear much te musculature cegment of the via a slitelike of this organ mucosa in the inthe oviduee ady to be laid. f che epithelial fas epithelium epithelium has v glands in the vesicular, and sum the rolti- Ih and then of Ek st columnar, cells Some ofthese cells are ciliated and slim, others non-iliated and more voluminows with indications of intense mucus secretion. The mucous membrane of che magnum contains numerous tightly packed, branched, tubular glands. During periods of high activity, these glands show ampulla-like dilations formed by theie mainly protein-containing secretion, Tm the region which borders on he isthmus, che previously single-layered epithelium becomes scratified, the number of ciliated cells increases and tle mucous membrane loses its glands. Inthe isthmus the secreting epithelium becomes lower and double-layered and che mucous membrane contains protein-secreing glands, though fewer in number than in the magnum. The last region ff the isthmus and the neat segment, the wlerus, are morphologically very similar both in expect of the covering epithelium and the glands. Flowever, the numerous glands of the werus produce 2 calcium-containing secretion. The mucous membrane of the vagina is aglandular and the ciliated epithelium is of columnar type ‘The avian egg: Before discussing the functions of the oviduct and its various segment, itis first necssary 10 describe the structure of the avian egg. The egg of birds consis of: 1.) The fertilised or unfertlised egg cell or sphere of yolk -which is produced by the ovary and already has a thin covering yolk membrane 79/a—c). 2.) The sphere of yolk is surrounded by three layers ‘of egg white or albumen (/g—i); these are of varying viscosity, the outer and inner layers being Jess viscous than the middle layer which later is of much greater volume. 3) The cbalaza (/f) are spirally ewised, cord-ike seuctaces which are suspended in che albumen and tightly attached t0 the yolk membrane. 4.) The double-layered shell membrane (/k) between which lis the air chamber (i), at the blune pole of the egg. 5.) The calcium sbell (/m) and on the outside the exticula which can be compared to a covering of varnish. The contribution made by each segment of the ‘oviduct tothe formation ofthe various components ofthe egg is discused in detail below. Fig.79. Diagram of «ben's ane whereof yolk doubleane SE a enbetns Fernie Sis, Seat gelee pbs e he Soler falas i eet sha Tice, fer sacs lope b vicoet Eyerstd fom ln ico yer 1 se dabets cleo shall wat ‘The following dasa on the chemical composition of the hen’s egg is derived from Mexnen and Raver (1958): In an egg weighing 58. the shell accounts for 6g, the albumen for 33 g and the yolk for 19 g. The yolk consists of 48:7 %o water, 16:6. protein (ovovitellin and livetin), 3246 Ya fat (wiglycerides of palmitic stearic and oleic acids, and also cholesterol, lecithin and cephalin), 1% carbohydrate and 1-1 /o minerals, The egg white consists of 87-90 water, 10°6 *s procein (ovalbumen, ovoglobulin, ovomucin and ovomucoid), 09% carbohydrate and 06*/e minerals. ‘The egg shell is made up of 16%. water, 33 %a protein and 95-1 %fy minerals. The yellow colovr- ing of the yolk is derived from dietary pigments, such as lutein, carotin and ovoflavin. Traces of the latter are also present in the egg white. The yolk and albumen also contain vitamins A, B 1, B2,D,E,F, Hand K. Egg formation: At the time of ovulation the follicle, which is ready to rupture, is placed into the correct position by active movement. It is then grasped by the muscular infundibulum and the sphere of yolk, enclosed in its soft yolk membrane, is liberated into the lumen of the infundibulum. ‘The yolk membrane, which was already formed in the ovary, is now surrounded by a concentrated,82 Sex organs rmucin-containing secretion in the form of a delicate fibre network, with the result that the screamlined yolk is loosely surrounded by a double-layered membrane. ‘This membrane is sezi- permeable so that, with the existing osmotic gradient, water can enter the yolk from the albumen By this means che weight of the yolk is made up tots ultimate value 40-5000 of the toral albumen mass is added in the magnum of the oviduct. The remaining 5060 is supplied by the isthmus and the uteres. The albumen (/g—i) supplied by the magnum is Ind down in more homogencous layers (/g, i), between which are denser structures (/2). The round sphere of yolk is thus surrounded by concentric lamellae of albumen. At this stage of the passage of the egg down the oviduct the albumen is not yet divided into the three viscous layers mentioned above, nor has the differentiation of the ohalaza (/f) taken place. On the other hand, the formation of the shell membrane (79/k) commences atthe isthmus end of the magnum, where the secretion is very mucin-rch, and is completed by the secretion ofthe isthamus. The fluid, provsin- containing part of this secretion provides the lest viscous outer layer of the albumen (/:) while the Keratin-containing component goes into the formation of che remaining part of the shell ‘membrane. This has a double-layered fibrous structure. Later, the outer layer combines with the calcium shell (/m) and the inner layer is in close contace with the albumen. Only after the ex has been laid docs the air chamber (ll) develop at the blunt end, berween the outer and inner la~ mellae of the shell membrane "The chalazae ((j) develop in the isthmus and uterus. The loosely applied yolk membrane intially projects into the albumen both towards the pointed and the blunt poles. Since the egg revolves as it travels down che oviduct, these projections of the yolk membrane, to which are Added fibrous components of the albumen, become spirally twisted because they remain firmly atached o the yolk. Theie other end, however, lies fre inthe albumen. During the development (of the chalazae, the less viscous albumen is liberated and forms a thin layer azound che yolk mass. Thus the yolk becomes suspended by the chalazae and it ean move within the egg, so that its germinal dise, situated at the animal pole, is always uppermost, while the heavier vegetative pole falls into the lower position Formation of the calcium shell (/m) commences as soon as the egg, loosely surrounded by its shell membrane, reaches the wide uterus. During the first phase of this process a watery, protein- containing component of the uterine glands pasies through the semipermeable shell gland into the albumen to make up its final volume. But apart from this the uterine secretion provides the calcium shell. The shell of the avian egg consists of a layer of calcium spheries (80) which a made up of radial aggregates of calcite crystals, The ste of formation (/b) of these shell constieuents is om the shell membrane (/a). These components are made up of the mammillary layer (Je, consisting of calote-ike structares embedded in the shell membrane, and the spongy layer which is composed of the inner cone-shaped (/e) and the outer columnar (/f) exospherits(/d) extending tothe shell sur- face. Although the exospherites touch, pore canals persist beeween them allowing gascour exchange with the en- vironment. The calcium shell measures about 0-32 mm in thickness and it contains aot only mineral but also small amount of organic matrix composed of a protein- polysaccharide complex. Coloured shells also contain protoporphyrin (ooporphyrin), a haemoglobin break- down product. Jn the fowi the egg requires about 24 hours to be completed and ready to lay but the time taken +0 pass through the various segments of the oviduce differs considerably. It remains inthe infundibulum for 0:33 hours, inthe magnum 3 hours, in the isthmus 117 hburs and in the uterus 19 hours. Daring oviposition the last part of the oviduct, he Fig. Two calcite sphesites of che ealcigm shell of CTE ge tw. stmnoss 180), . iT exoshecte cones feslisats growt linesresult that the om the albumen. ‘The remaining by the magnum revures (/A)- The this stage of the cee viscous layers the other hand, magnum, where be fluid, provein- sumen (/i) while vart of the shell mbines with the aly after the exg ter and inner [a yolk membrane es, Since the egg rnc, to which are yy remain firmly the development around the yolk ‘the egg, s0 that wrrounded by its watery, protein- Shell gland into son provides the = (80) which are shell constituents illary layer (i), ongy layer which a) and the outer 410 the shell sur- sore canals persist age With the en- about 032 mm sineral but also 2 ved of a protein- ells also contain rmoglobin break 24 hours to be sme taken to pass 2 oviduct differs afundibulum for in the isthmus the oviduct, the calcium shell of orlset) 2c mmm layer Embryonic membranes 83 vagina (75/b; 77/0) is projected into the cloaca and then everted along with the cloaca, so that the finished egg is transported to the exterior directly from the uterus, The outer euticle covering. the egg is sticky when ic is first laid but chis dries rapidly. Ic is not uncommon to encounter eggs which are abnormal in size, shape or content. Such are the large double-yolked eggs. Both yolks may have originated from one follicle, in which case they have a single yolk membrane, but it is more common for two independent follicles to ovulate ‘within a short period of each other and in such cases each of the yolks has its own membrane, Dwarf eggs ate small eggs which either contain only albumen or only yolk. Pathological conditions of the oviduct lead to the development of kidney-shaped or spindle shaped eggs, which contain coagulated protein or fluid albumen. Malfunction of the uterus causes deficient development or ‘complete absence of the shell. Finally ic should be mentioned that foreign bodies, oz even parasites, entering the oviduct from the cloaca by antiperistaltic movement may also give rise to abnormal cass Development and function of the embryonic membranes in birds Te has already been explained thatthe female sex organs of most bigds, and all domestic species, consis in the postembryonic site of only s left ovary and the let oviduct, the later being divided into various ‘orphologically and funetionlly dnine regions. There organs are under hormonal control and only reach their fall development during the reproductive pesiod aftr the onset of sexual matorty, The product of {hese organs i, OF course, the aun egg, which fea closed system in whidh the embryo develops aul the Bide is ready 0 hath ‘The development of the zygote begins upon ferlisation of the mature ovum inside the female generative tcact Te goes through early developmen: (Ulastogensss) to the formation Of the blstorern THe dencioy imental process is temporacly interrupted by the deposition of the egg; It contnucs to completion only here flee of hu rng brooding Bro ae seco overepuron ania, An von oe Sulsite for this ype of teproductt is thatthe egg must contain all toe nutientsrequced By the eer Yah the excepsion of oxygen, whichis derived from the Stmosphere. It must also powess 4 thell whi protects it from ‘watelom and mechanical damage. The bird receives some of these substances irom the vary since the polylecithal ovum i provided with a large amount of yolk: Tey released inthis condition ‘rom she ovary and is received by the oviduct whees the albumen, conssung of protein abd much wate, sd, as walla the shell membrane and the calcium sel Birds age apnione verebrat,dhractrised by she face that only a portion of the germinal material present after fertilisation is used for the actual development of the embryo, while the sminder forms the niyo marron Te pects have she futon of pret Ue cae tne medal iniuey, of supplying i withthe notrientsconeained inthe eg, of removing is break down products ad of felting gaseous exchange. By means of 4 complicated procesy the amnion (G0.a/3) develops fom te cuter and the midale germinal layer (ectoderm and parietal meroderm). The snnion ipa lui flled vesicle Sn hid she floating embryo (/1) can develop. Ancther membranous organ, the yolk sat U3), are fom fhe inner and middle germinal layer (ovtoderm and slvceral mesoderm and te ramains connected ta the Jntesnal trac through a salle. THe yolk sae surrounds the sphere of yolk and develops «systeat of blood ‘sels, the yolk sac citculaion. The min veils of this system are thea and ov, ompbalomesentericae ‘hic join the blood cizeulatory system of the embryo ehrough the navel ‘The mations contained ia the Yelle (6) are pecpared by the cells of the yolk ste for sobueguent metabolism and easel tp the embryo Ey the vessels of the yolk sac.‘Thue the yolk sa. sontioually decreases tn size uot afer harap 9 ‘ken through the navel into the body eavity aod she gut. Ite remaining contents are ulised durieg the First days of the cide’ independent existence. Simoltancoualy with the Tormasion of the amnion ancthet smembrane, the cborion, also develops from ccoblase and parital mesoblast. Te surrounds the yolk sac tnd ‘fen buen tpt rm hom by gap the Cele ll) whe a lan of the clones, grows through the navel opening into the exocos, ‘The wall of the allanteis is derived from rode it burl moder Te panne eoanecon wih he donee rough th lane dat nd by this route, ie collects the ule acid-ridh embryonic urine whichis initally sereted by the pronephos tnd later by he metanephros, Thar the allantos dilates tt0 a "ating sac” whith may te likened to sm Nembryonic urinary blader” (9). Av s rere of this dlation, the outer surface of the allantois contacts the inger surface of the chorion and these two fuse to form the allaatodiorion (7). At the same me at ‘he allacos is being formed, omyphale arveties and veins prow oot of the embryonic ceculatory stem $0 ‘tscularize the allsntodhorion. Ia consequence there fr Sow second extraembryonic evclavon, te Sllentoic cirenlation, which underees gaseous exchange. This can only be achieved, however, if contact Bade herve he lncothoron an fe sl membrane (1) The porous sl (1) meek allows the trans of oxygen ard carbon dloside but it also provides the embryo wish calcium salts: Recent Investigations have show that the egy albumen (s nov casted to the embryo by the slants cecalston, [ye llance ci il wih nee rcp wiened psphecally andthe Cone ino clave apponion ‘withthe amnion and yolk sae Together with the chorion ie forms the so-called albumen sae (10) which Ccoizs the alloumen, now concentrated chrovgh lors of swat tothe yolle The conics between simaion fad chorion makes posible, at a later sage of development, for the albumen sac to break rough {nto the amoiotie exvity (1). The albumen pushes ehrough whe canal (10) Fonned by this roptare mines ‘with the smniotc fai, and tx swallowed bythe bird and gered.Fg tos, Longigadina on the tik day of ine (eer aN Racost, 1961 4 entero: 2 wma consising S yolk cs 6 yolk 7 allanoe rig filed with embryonic Mbemen; 20" abemeo, etal, 7 Sul tbeane 1 ee sell Sih pres ais aber ‘The allantoic circulation atrophies even before the developmental period is completed. The chick perforates the embryonic membranes and the shell membrane with its egg tooth and geinr access 20 the Ealacged sir chamber (113); t Begins to breath. In the fow! this takes place two to three days before hhatdhing and chirping noises can be heard during this time. Finally, and again with the aid ofits egg tooth and its limbs, the young bied breaks through the shell and Jeaves she egg. This occurs in the fow! alter an incubation period of 20—21 days, im the duck 26—28 days, in the goose 2833. days, in the rurkey 2630 days and in the pigcon 1316 days ‘The fowl dack, turkey and goose are autophegous, tha isto say, they leave che nest immediately. At hatching they have complete down plumage, Theis locomocor apparatus and other organ system, including the sense organs and nervous system, are suficienly developed to perform all the necessary fonctions for their survival and further independent development. Parental care is confined to help in finding food, to protection agsinst dangers and ro providing warmth during slep or cold westher. Pigeons, on the other and; are tusttorial birds At the Time of hatching all the organ systems, except the digestive, and Feary paras are gpl developed inch newly Bred abn As fa they are ake, nd intensive brooding and care before they become fledgedoesigedial ch'day of nt ed, The hide se acu the ree days belore of ies sag tooth se fowl akcer 20 vin the carkey Tee Sue Circulatory System Blood and blood circulation ‘The blood consists of the blood plasma and the blood cells, ‘The plasma is a viscous, aqueous solutions of various proteins such as albumen, globulin and fibrinogen, of carbohydrates and salts and it also contains fats and lipoids in fine suspension. These substances, as well as hormones, vitamins and enzymes which may also be present in blood, are all necessary to maintain the body cells. But the plasma also cakes up, for ultimate elimination from the body, break-down products resulting from metabolic processes; such products include carbon dioxide, uric acid and other nitrogen-containing compounds. The plasma has the ability to coagulate blood. This is an enzymic reaction requiring the participation of special blood cells, the thromboeytes, by which the fibrinogen of the plasma is changed in a series of steps to fibrin, Together with the blood cells, it forms the red blood clot. The fluid part of the plasma, the yellowish, clear serum, is expressed during formation of the clot The cellular components of the blood are 1.) the red blood cells or erythrocytes, 2.) the white blood cells or leucocytes and 3.) the thrombocytes which in birds are true cells, sometimes called “spindle cells” because of their shape (ef. table and colour plate 111) In birds the erythrocytes (A/1) are nucleated, oval cells. They contain the blood pigment which is the iron-containing h2emoglobin; chs substance enables the erythrocytes to carry oxygen. ‘The white blood cells include the polymorphonuclear granulocytes, the lymphocytes and the monocytes. Average numbers of blood cells in domestic birds? Erythro- Leucocytes piearbe, a ! 64-25, 21-05 575 3455, aa a 24 |60:2—68-4) (1924-2) | (5464) | (46) G—#2) i 28 | o45 | 4920 | 305 | tase | tas oss | Pigeon |245—99/99—723 (49-2—60-6) (29-4332) (58-66) | (226) | 18-3) ee After Scusnuse, 1950/54 and Sounean, 1958,86 Circulatory system ‘The leucocytes are capable of amoeboid movement and can leave the intact blood vessels. Daring this process, known as leucodiapedesis, the leucocytes slip through the thin capillary wall into the surrounding tissues at the same time considerably modifying the shape of their cytoplasm and nucleus. A property of the granulocytes is that of phagocytosis; they are attracted to sites of tissue damage or to accumulations of disease-producing organisms by chemotaxis and they then surround and ingest cell fragments or bacteria which they bréak down by enzymic action. "The granulocytes are nearly spherical in shape and they have a polymorphic nucleus. ‘Their cytoplasm contains granules which react differently co various dyes and it is possible, by using appropriate stains, to subdivide this class of leucocytes. By far the greacest number contain rod~ shaped, acidophilic or eosinophilic granules and they are known as psewdoeosinopbils or heterophil granslocytes (Al6; BIS). A smaller number of these white blood cells contain round eosinophilic granules and these are the eosinophil granulocytes (A/7; B/3). A third, numerically very small, sroup ate the basophil granulocytes (A/8; B/4), so-called because of the selective affinity of their sgransles to basie dye. ‘The avian monocytes (AIA) are difficult to differentiase from the large lymphocyzes (infra), because they are similar in shape and size. However, unlike the round nucleus of the large Iymphocytes, the nucleus of the monocytes shows an irregular indentation, so that it sometimes appears kidney-shaped. With the Romanowsky stains their cytoplasm shows a delicate bluish-geey ‘The Iymphocytes are far more numerous than any of the other white blood cells. Alshough similar in shape and staining reactions, large and small varieties of lymphocytes are differentiated (13, 25 Bi). Blood coagulation is initiated by the break-down of the thrombocytes (A/5; B/2) which, unlike the functionally homologous thrombocytes or blood platelets of mammals, are true nucleated dls. Blood formation. The first blood cells, and also che endothelial cell lining the blood vessels arise from smerencynal cements called snioblats which are stoned in the blood islands ofthe area easculose of the vninal vesicle apd the wall of che yolk sac. Other sites of blood formation during development are the {Retr pees and thyrnus bur the bone marrow finally becomes the organ of blood formation tn the higher varebrates, All types of blood cell including Iympboeyees, are found in various stages of maturation in The bone marrow, However, lymphocyte production in birds also takes place in all types of lymphatic ‘ge, including the lymph nodes tn dudks and gese, "Fh faneoa of the Bial aze numerous a iciadesoplyng te due apd calla with bane vit for their maintenance and function, Among these funetions can be counted the transport to the tissues of intel om edger og sorageortans snd of hormones produced bythe sarin gas, nd the Supoly of oxygen ffom the lange. Equally important is the removal from the tissues co the site of SiREnacon of Larabolc products nigh ay carbon dioxide and aitogen-contsining breakdown product of proteins, The blood muse algo carry so the storage organs certain substances which are present in excess of Ereabolic requirements while ye ¢ further function i regulation of water and electrolyte metabolism and Iintenance of a conmant body temperature. ‘Blood circulation. In order to accomplish al ts Functions, the blood has to be maintained in circolation through 2 closed system of tubes, the Blood vesels by the action of the hear. The left veniile expels Guygenned blood through the covts aad thas round the body. The arteries, arising from the sorta, cay fa crery organ through brandhcs of ever decreasing diameter snd finally, having fist become arterioles, end In a dence network of capillaries, Thee one to form ventle which continue a veins and end in the single nuded and double crane! cena cava: In this way the Blood ix carried t the right atvinm and then the right Stntvices This pare of the blood cireuc is known aa the greater of systemic cremation. The venous, o ‘SZirypenated, blood is eared by the small or pulmonary Gienfation from the right ventricle through the pubpotery arteries to the lang, whence the arterial blood, now oxygenated, is carried through the ‘uimomaryceing to the lef atrium and then the left centile, thus sanding the systemic circulation anew. ‘Serucrure and function of blood vessels. The wall of an artery is,made up of thrce layers: an inner layers the intima, a middle layer the media, snd an external coat, the adventitia. ‘Three togions can be iliefentated in the system of arteries originating from the heart The arteries nearest the heart are of the liste type, in which the media conssts mainly of elasie elements and-whidy have the property, of feverble distension, For this reson they are ablg to “nore” energy and thot ating ike an air camber, {hey ean vanslate the surges of Blood coming from the heart into a continuous steam, The more distal [ntti act as dstribucors of te blood to the various organs. The media of these veel & made up mainly ‘Sf smooth muscle which is under the contol of wasomotor nerves By alternate dilatation and constriction Gf the lutea they regulate the pressre gradient in the arterial side of the circlaton and chus control, Scoring tothe fonetonal requirement of the later, the amount of blood passed vo the organs. However, helo soppy to the organs can be controled mach more effecvely by the ererols and he capllaces frising from chem, than by the lagge arteries. The arterioles are able to contract to such an extent that fhe lamina can be completely occluded. Acting like a valve, they can temporarily reduce the blood flowblood vessels. capillary wall teir cytoplasm ted to sites of ind they then sucleus. ‘Their ible, by using contain rod- + of heterophit 4 eosinophilic iy very small, finity of their cytes (infra), fof the large tit sometimes ate bluish-grey ells, Although differentiated whi, unlike crue nucleated sae from Sci fe snip the higher Possess Sof lmmphate othe dius of ‘organs, and the wo the ae of vn product of Sie exces of ‘metabolism sod 4 in deatation Sire pe, Se ee tee Aon fe Seeoraily. layer: an nner regions can be jesteae a de fn tr camber ‘The more dial made up mainly Sn comnieion id chur contol, pone, However ihe capaci an este shat blood flow PLATE IIL A. Blood celts of « fowl 1 eythreyee: 2 aall mphoryte; 3 lage Iyaphoce; 4 monoye: 3 thrombocye; 6 pteuotsnopil granulocyte (hewrophi) T cone gratlocr# boop praslacre 5. Blood cells of the fow!. (Afr Kasuume sed Gase, 160) 1 small and lee Iymphoryer 2 shrmbocys se; # basophil granulocyte; §pueadoevnopilHeart 87 to a relatively jnactive organ and consequently provide preferential supply to an active organ. The [pillres, the lumina of which may be 9 narton? shat individal blood ells cam only jst pest through, Sre alone responsible for mediating the exchange of substances ia both directions from the blood to the {fnercelllar Hui. The counties numbers of capillaries whose walls consist only of a single, thin layer of dothelum and a bas membrane, representa vast enlargement of the vasclar system through ‘which the Ela flows slgitely slow'y. Fhe rec of the blood tothe hear is va the venows side ofthe cireaation ‘The capillaries join vo form the posteaptllary veins or venules and these in turn unite into the larger veins, which later gencrally follow the Coarse of the artic. ‘Thes lumen is generally wider than thar of che accompanying artericy bat ther media, whichis made up largely of connective tinue i thinner. Venovs blond. under low presace and its return flow to the heart is facileated by haemodynamieally acsive structures in the veins Ties inclade aloer which prevent the backflow of blond. But she venous blood also rectives active Bro- pulion through the so-called svtentavenons anastomoren, which connect small arteries 0 Wein an6 49 Rorecicate the capillaries. The hacmodynamie efect of those anastomones, which can be opened or closed required, lie in che face that the arterial blood under higher pressure can be shunted into the veins ‘Then the oyotem is ope, oo that the low presure venous blood i propelled towards the hess, Blood-vascular system Heart (cor, cardia) (43, 44, 45, 46, 74, 76, 77, 8184) ‘The heart of birds is relaively large (43/o; 44/1; 45/n; 46/k; 761d 77/f, ). Tt is enclosed by the pericardium and lies within the cranial part of the cavum eardioabdominale cf. p. 39). Ts long axis, being slightly deflected so the right of the midline, runs from craniodorsal to caudo- ventral. The base of the heart is directed craniodorsally and lies ac the level of the second rib, while the apex of the heart points towards the sternum and is situated on a transverse plane passing between the fifth and sixth ribs. Lying dorsally against the base of the heare are the lungs covered by the cranial segments of the horizontal septum, the two primary bronchi, the sysinx and the oesophagus. ‘The dorsal surface (facie atvialis) and the lateral surfaces of the heart are embedded in the liver, whichis enclosed in its peritoneal sac. On the left side the heart contacts the proventriculus. Its ventral surface (facies auriculars) is sivuated cowards the sternum. Finally the heart, or more correctly the pericardium, is clotely related to the diverticula of the cereal, clavicular and eranial thoracic ‘The outer fibrous layer of the very thin pericardium is connected by delicate strands of connective tissue to the hepatoperitoneal sac and the above-mentioned air sac. Itis also Fused with the horizontal septum where it is closely applied cranially and laeerally, and with the oblique septum (fp. 38). The fibrosa of the pericardium continues basally into the adventitia of the large vessels and, through the li. sternopericardiacum, to the sternum. ‘The internal lining of the cavnm pericardii consists of the lamina parietalis, a serous membrane fused with the fibrosa. At the base of the hearr it surrounds the origin of the large vessels where itis reflected onto the heart as the epicagdinm or lamina visceralis. The pericardial cavity contains afew drops of serous fluid, the liquor pericardi. ‘The dark red to bluish-red coloured conical heart (cor) of the bird consists of epicardium, myocardium and endocardium. Because of the high basal and energy metabolism of birds, which have a body temperature of 40—42° C, and a very rapid heare rate (200—300 heart beats per minute in the fowl and 5C0—800 in the sparrow), the heart has a correspondingly great relative weight. In the fowl it accounts for 0'5 ro 1-42 / of the body weight, the greater figures applying88 Blood-vareui system to smaller breeds. In the turkey it accounts for 05 0/s, in the duck and goose for 0-8 ®/e and in the pigeon, which is a good flier, for 1-1—1-4 %a of the body weight. "The avian heart has a slightly flattened dorsal surface (facies atrialis) (81), a ventral surface (facies awricularis) (82), a rounded, slightly convex left (81/1) and a somewhat concave right surface (/2). As in mammals, it consists of a venous right and an arterial left half, each with an atrium and ventricle. A distinct coronary groove (sulcus coronarins) (/3) filled with adipose tissue ‘separates the atria from the ventricles lying below them. The longitudinal grooves (sulcus inter- ventricularis paraconalis (82/1) et subsinosus (81/4) are insignificant. The two atria (81/5, 6) rise cupola-like above the coronary groove and surround with their ventrally directed auricles (axri- ula cordis) (84/1, 2) the aorta (82/16; 83/12; 84/3) and the truncus pulmonalis (81/15; 82/135 83/113 84/4) ‘The right atrium (ateium dextrum) (74/c’; 81/6; 82/2; 83/1) has a greater volume than the left. Its wall, which in some parts i exceedingly thin, is supported by muscle bundles (mm. pectinati) (8213), which are arranged in net-like fashion and project into the atrial lumen. Tt is Separated from the left atrium by a thin septum interatiale. Duving embryonic development this Septum has sieverlike perforations which allow the blood to pass from the right +o the left erium, These slitlike openings disappear without tace with the onset of pulmonary respiration. ‘The three vende cavae terminate in che facies atrials. The very large cava cand. (81/75 83/2) enters the basal pare of the right atrium chrough a roundish orifice. In the bird there are two to. cavae cranm the right (81/8; 83/3) enters the atrium above the termination of the caudal vena ava, while the left cranial vena cava (81/95 83/4) enters the right atrium immediately above the Coronary groove and to the left of the caudal vena cava. Here also terminates the v. cordis media (83/3) which runs on the dorsal surface of the heart. The v. cordis magna (84/5), on the other land, runs along the ventral surface over the base of the heart, between the pulmonary trunk and the left auricle, to terminate in te right ventricle opposite to the entry of the caudal vena cava 'A thick muscular trunk (82/4) projects into che lumen from the roof of the right atrium and from this issue smaller, interconnecting bundles of muscle tissue, the mins. pectinati. A stout roscular ridge forms a connection between the orifices of the three venae cavae. This is che zm, pectinatus valonlaris which arises from the muscular trunk (82/4) and runs under the opening of the v. cava eran. dex, (82/5), chen forms a semicircular fold around the right border of she orifice of the v. cava caud. (6). As a muscular bridge it then forms the dorsal border of the left cranial vena caval orifice (/7) and continues as a second tall fold. Both these muscular folds, specially the very tall right fold, act at valves which close the caudal vena cava (/8,9). But be- Catse of ther relationship to the oxtia of the two cranial venae cavae, they are also able to prevent the backflow of blood into these veins. On the floor of the right atrium there is the moon-shaped ostium atrioventriculare dext.(/10), this shape corresponding to that of the right ventricle. The medial border of the ostium is formed by the interventricular septum which projects into the atrium, while the lateral border is formed by a curved muscular ridge lying against the septum. “The left atrium (atrium sinistrum) (74/c; 81/3; 83/6) is less voluminous than the right but it has a thicker wall, Te to0 is supported by the muscular trunk which continues from the right atrium through the interacrial septum onto the roof of the left acrium where it also divides into numerous mm. pectinati, disseminating along its wall. The ostixm avvioventriculare sin., surrounded by a ircular muscle ridge, is in the form of a funnel-shaped depression on the floor of the left atrium, The two pulmonary veins (81/10, 11; 83/8, 9) terminate on the facies atrialis in a diverticulum of the left atrium. ‘This secondary chamber is separated from the main cavity of the atrium by 1 sickle-shaped mascular valve, suspended from the roof of the atrium, which prevents the back- flow of blood into the pulmonary veins. In the avian heart the musculature of the atria is completely separated from that of the ‘ventricles by fibrous rings, the anuli fibroit (the skeleton of the heart). These are particularly well developed at the origin of the aorta and pulmonary artery as well as at the right atrioventricular frifice. Cartilagenous supporting structures, the so-called cardiac cartilage, are present in the fibrous rings around the origins of che aorta and the pulmonary arteries. ‘The right ventricle (wentriculus dexter) (74/6; 81/12; 82/11; 83/7; 84/6) commences at the facies anricularis of the heart, continues beyond its border to the facies atrialis and lies against theHeart 89 80/9 and in the ventral sueface ‘concave right 4 each with an B adipose tissue ss (suleus inter- ‘ia (81/5, 6) rise L auricles (auri- (BU15; 82/13; ‘lume than the > bundles (mm. ial lumen. Te is evelopment this ight to the left i tary respiration. ud. (81/7; 83/2) A there are two the caudal vena iately above the ‘v. cordis media Fig, Mesee of the earker, Doct Heart of the verkey with tighe 5), on the other ' ibe, Tae ei Bilt Td Vedieiele, spend, Vente, puary tema one fa? Slew lester bon 1 aeusintrrenscelari parnrmlity 2 sei dees vena cava, Plenty 8 aun does Pe cart 3D mn pevtinny ¢ munulat tandie 9 oie of the ighe atrium and Siirbo o polmoaly sins 17 pulmonale FET eae en tes Pht ted 3 ile vave of ctinati. A scout Fa cael: 10 com seloventeace deze 1 ve vac. This is the ) scene) les der the opening ‘ext 18 aorta; 19, 3%, bradhocepbalica Ca ate near 17218 «palate tin ad eee nt border of the a ood be BT Seinen saan ene 320 3° order of the left ale aorta dese muscular folds, 1 (8,3). But be: lefe ventrile in the form of a slit-shaped pouch. Tt makes a shallow curve towards the point of > able to prevent the heart but only extends about two thirds of the way to apex of the left ventricle "The septum interventriculare bulges far into the lumen of the right ventricle (74/b"; 82/12) dare dest, (10). which, in erose-seccion, appears asa sickle-shaped cleft. The outer wall of the ventricle bears low ostium is formed muscular ridges which are distinct in the region around the origin of the a. pulmonalis bur sorder is formed gradually disappear towards the apex. Ia the region of the slt-like right atrioventricular orifice (82/10), the musculature of the external wall is reflected into the lumen of the ventricle, thus giving rise vo a triangular muscle place (74/b'"'; 82/21). The atrioventricular orifice is bounded laterally by the chickened border of this reflection, which commences at the ostium: pulmonale to descend along the atrial contour of the ventricle. ‘The sharp free border of the muscle plate, running parallel to its insertion, lier close to the bulging interventricular seprum. It is also fixed to the roof of the ventricle by means of a muscle bundle (82/21") and to the septum by a sail-like membrane (/21”). This muscle plate, situated at the site of entry into the right ventricle, is particularly characteristic of the avian heart. It acts as 2 valve for the right atrioventricular orifice preventing the flow of blood into che atrium during ventricular systole and is chus analgous : to the tricuspid valve of the mammalian heart. fo the left of the atrioventricular valve is the outflow region of the right ventricle and here, rising between the two auricles, is the conus arteriosus and the ostium sranci pulmpnalis (82/13). Here is situated the valva trunci pulmonalis (14) which consists of three valoulae semilunares so arranged that one is to the left and the other to the right of the ostium and the third is on the interventricular septum. These valves prevent the : backflow of blood during diastole. ‘The left ventricle (ventriculus sinister) (74/6; 81/14; 82/15; 83/10; 84/7) is in the form of 2 ‘ommences at the thick-walled, hollow cone which in cross-section appears almost round. The outer wall and the id lies against the interventricular septum (74/6’) are about three to four times thicker than the wall-of the right the right bur ie , the right atrium surrounded by a the left atrium. a diverticulum of the atrium by tevents the back- rom that of che particularly well {atrioventricular e present in the90 Blood-vasculat system ventricle. The left ventricular aspect of the septum is smooth but the ourer wall of the left ventricle bears muscular ridges of varying height which protrude into the lumen. From the ansins fibrosus surcounding the ostium atrioventriculare atises the walva atrioventriculare sin. This valve, consisting of a double layer of endocardium and intermediate fibrous tissue, is extremely thin and almost transparent. The free border of this tricuspid valve carries a varying number of chordae tendineae. One cusp is situated against the septum while the other two cusps are on the outer wall. One papillary muscle is allocated vo each cusp. They vary in appearance and may be subdivided into individual muscle bundles. The chordae tendinege are attached to these papillary muscles, one of which originates from che outer wall, the second from the septum while the third ‘originates from both septum and outer wall. The left atrioventricular valve prevents the backflow of blood into the atrium during ventricular systole. ‘The aorta (82/16; 83/12; 84/3) originates from the roof of the left ventricle between the right auricle and the conus arteriosus of the pulmonary trank (82/13). The aortic valve (valoa aortae) js situated in the ostium aortae and, like the pulmonary valve, it consists of three valeulae semi- lunares. Blood supply to the heart In birds the heart is supplied with arterial blood through the two coronary arteries. ‘The a. coronaria dext. (84/8) arises from the sinus of the right semilunar valve of the aorta and after bbut a short course it divides into a ramus supf. (84/9; 83/13) and a ramus prof. (84/10; 83/14). The superficial branch runs in the coronary groove to the right and, reaching the facies atrialis, it gives off branches to the wall of the right atrium and ventricle. The lacter branches to the ventricle {quickly enter the depth of the musculature where they ramily furcher. ‘The very chids deep branch runs in the interventricular septum towards the apex of the heart and divides into ewo large branches. The subdivisions of these leave the septum and enter the ‘musculature of the left ventricle from ventral and dorsal. They anastomose on the left border of the heart. The right coronary artery supplies, via its deep and superficial branches, a very large part of the heart, including the wall of the right atrium, a large part of the sepcam and the wall of the left ventricle, ‘The a. coronaria sin. (84/11) acises from the sinus of che left semilunar valve of the aorta. Inicially its course is directed to the base of the heart between the pulmonary trunk and the left atrium, It then divides into a ramus sup}. (84/12; 83/15) and ramus prof. (84/13). Below the left auricle the superficial branch enters the coronary groove and runs along it to reach the facies atrialis. The thicker, deep branch enters the interventricular septum where it gives off secondary branches to the wall of the left ventricle. These branches, as well as those of che superficial branch, anastomose on the jacies atvialis with secondary branches from the ramus prof. of the right coronary artery. Thus the left coronary artery supplies the wall of the left and parts of the right atrium, a part of the septum and parts of the right and lefe ventricles. ‘Two veins with wide lumina and several smaller veins (83/16) return the blood from the cardiac circulation to the right atrium. The wide o. cordis media (83/5) ascends in the sulcus subsinosus to the right atrium where it verminates immediately above the coronary groove. The vvein collects blood which had originated mainly from the deep branch to the right coronary artery A second large cardiac vein, the v. cordis magna (84/5), originates at the apex of the heart and runs along the left ventricle enter the coronary groove whence it proceeds, under cover of the pulmonary trunk, to the right atrium. It enters the right atrium opposite the ostium of the caudal ‘vena cava, This vein collects blood from areas supplied by both the left coronary artery as well as by the deep branch of the right. As well as numerous small veins which enter the right ventricle direct, there is a larger vein running along the right border of the atrium, which conveys blood from the superficial branch of the right coronary artery, and another which collects blood from the areas supplied by the superficial branch of the left coronary artery. ‘Conducting system of the heart ‘The conducting system of the avian heart consist of the sinoatrial node, che atrioven- tricular node and the atrioventricular bundle. The nodus sinuatrialis (85/1) is siveatedwall of the left From the anulus ieulare sin. This sue, is extremely ying number of cesps are on the ance and may be o these papillary a while the chied ants the backflow vetween the right ze (vals aortae) ve valoulae semi- tay arteries. The aorta and after 4/103 83/14). The 4 atrialis, i gives to the ventricle pex of the heart m and enter the he left border of hes, a very large tum and the wall ‘ve of the aorta tank and the left 1), Below the left teach the facies 1s off secondary sperficial branch, ‘of. of the right and parts of the blood from the ids in the sulcus ary groove. The 2 right coronary of the heart and der cover of the am of the caudal y artery a8 well xe right ventricle 4h conveys blood s blood from the 3 the atrioven- 5/1) is situated Heart Bea ventricles with che large vessels aad cardiae ilved supply of the fowl Don vse, fate avin Fig, Plassoid corrorion preparation af red yencricien sith the. cacline blood sop {he owl Vantal view Lhe 6,17 ae bradicephaliea ts and est 189, cave cnn, de cs bi, Poi | (sher Banas, i) 1 nodes sinotaits 2 nodes acoyensiaaias 5 daniels nstveniclce wth Fert ted"ins Sanding Brand of the ere teres bead of he ee ti* 92 Aneries in the wall of the right atrium at the site of entry of the caudal and left cranial vvenae cavae. The nodus atrioventricularis 2) is oval in shape and from i originates the fascias atrioventricularis (bundle of His) (/3) which euns through the amulus fibroms, enters the intr- ventricular septum and, after a short course, divides into right and lefe limbs. The right limb {eras dext. () rons below the epicardium towards the apex where it divides ro connect with the network of Purkinje fibres. Shortly after its origin it gives off a-branch which again ascends into the muscular right atrioventricular valve (/3). ‘The left limb (crus. sin.) of the fasciculus atvio- ventrcalaris (14) runs, also below the endocardium, on the left side of the septum to the apex of the heart where it also connects with the Purkinje fibres. Shortly after is origin from the bundle it gives off the socalled recurrent branch (/6). This ascends in the septum, enters the ainulus fibrovs, encizcles the base of the aorta from the left and connects with the bundle of Purkinje fibres which arise from the atrioventriculae node and run along the edge of the right atrioventricular orifice. Arteries In birds as in mammals one differentiates between the so-called small or pulmonary circulation, which carries venous blood from the right ventricle through the pulmonary trunk, and the so-called large or systemic circulation whose main vessel, the aorta, arises from the left ventricle and car~ ries arterial blood. Pulmonary arteries ‘The truneus pulmonalis (81/15; 82/13) arises from the facies auricularis of the right ventricle ‘where tis flanked by the auricles. Its dilated rooe is known asthe conus arterioss, The semibunar valves in the pulmonary ostium have been described in the section dealing with the hears. The short pulmonary trunk divides into the a. pulmonalis sin. and dext. (81/16, 17; 82/17, 18; 84/14, 15; 8612, 3; 87/1, 2). The lee artery crosses the left cranial vena cava (81/9; 8217) in a cranially convex curve and enters the left lung ventally, together with the primary bronchus and the pulmonary vein, Within the lung its branches (86/2) follow the subdivisions of the bronchi to terminate in the respiratory capillary network inthe wall of the parabronchi. The right polmonary artery crosses the aorta, which lies to the right of its origin, to gain the hilus ofthe right lang, Ts subsequent course i similar to that of the left pulmonary artery. Aorta ‘The aorta (81/18; 82/16; 83/12; 84/3; 86/4; 87/3) arises as the aorta ascendent from the left ‘ventricle. Close to its origin it has an expansion, the budbus. At the base of the heart itis surrounded by the left and right atria, the right auricle and the pulmonary trunk. At the level of its semilunar valve (86/5) it gives off the left and right coronary arteries (84/11, 8). Bending to the right it forms the arcus aortze and gives off first the a. brachiocephalica sin. and immediately afterwards the a. byachiocephalica dext. (81, 82/19, 20; 84/16, 17; 86/6, 7; 87/4, 3). As a resale it decreases considerably in diameter and, known now as the aorta descendens, courses berween the right lung and the oesophagus to reach the vertebral column at about the level of the 4th to 5th thoracic vertebrae. Between the two lungs, and accompanied dorsally by the oesophagus, it becomes the sorta thoraciea (86/4") and subsequently the aorta abdominalis(86/4"”). Along its caudally directed course it gives off the paired and single arteries of varying diameter, which will be described below in detail. Iv finally terminates on the ventral surface of the coccygeal vertebrae as the slender 4. sacralis media (86/8; 91/1"). Arteries cranial to the heart ‘The head, neck, wing and pectoral regions are supplied by the aa. brachiocephaicae (86/6, 7: 87/4, 3). Tehas already been sated thatthe left and right brachiocephalic arr ae, on ace the ether, from the aorta ac the base of the heart before the aortic arch bends to che Fight in craniodoral convex curve. Both arteries bend gently towards ther sie of the body, each crosing the primary brondius to teach the region of the shoulder join. Each brachiocepbatc artery now vides nt respectively the cavots com. dext et in, (869; 8716; 88/1) and ac subclvin deste,ad left cranial tes the fasciculus centers the inter= ‘The right limb ‘connect with the sain ascends into fasciculus atrio- ‘tum to the apex origin from the rum, enters the hy the bundle of Age of che right nary circulation, and the so-called entricle and car- e right ventricle «The semilunar a the heart. The 17; 82/17, 18; 81/9; 82/7) in a uy brondhus and of che bronchi to tight pulmonary ae right lung. Iss rns from the lefe ‘itis surrounded of its semilunar 0 the right it ftely aferwards osule it decreases inthe right lung to 5th thoracie + it becomes the ‘audally directed described below eas the slender alicae (8616, 7; s arise, one after > the right in its Ay, each crossing nalic artery now bclavia dext. et Arteries cranial co the heart 93 sin, (86110; 87/7). At the point of division a smaller artery arises and proceeds towards the caudal end ofthe trachea to send branches to the syrinx and the crop. ‘The a, carotis com. dext. et sin. converge and run cephalad, close co one another, along the ventral surface of the bodies of the cervical vertebrae and becween their paiced ventral processes. ‘They are covered by the m. longus coll. Shortly after they originate, each of the two carotid arteries gives off several branches, including the a, thyreoidea cran, et cand. (9816, 8) to the nearby thyroid gland and the small a. bronchalis to the bronchi. The a. oesophagica ascendens also arises here and it sends branches to the crop, teachea and oesophagus. To accomplish this, the left artery changes over to the right side under the trachea, and then takes a course berween the latter and the oesophagus (87/9). A thicker branch of the common carotid artery isthe a. vertebralis, che main derivative of whic, the a. vertebralis ascendens (86/11), enters the transverse canal of the cervical vertebrae. From there ie sends segmental vessels to the cervical musculature and, through the intervertebral foramina, to the spinal cord and its meninges. Finally, at the second cervical vertebra, it anastomoses with branch of the a. occipitals (88/3), which runs towards it. The thinner, caudal branch, the a. verte- bralis descendens, passes on each side between the tuberosity and head of the ribs up to the 4th or 5th thoracie vertebra, giving off the aa. intercostales and branches to the back muscles and to the spinal cord ofthis region. ‘The a. comes vagi (86/12; 87/8) also springs from che common carotid artery. Te accompanies the vagus nerve (87/43), Iying against che jugular vein (86/54; 87/27), supplies branches to the surrounding tissues and the skin and, in the upper third of the neck, subdivides into smaller arteries which may anastomose with similar vessels from the a. ocipitalis The two carotids diverge near the head and fork into an a. carotis ext. et int. (88/4, 5) at the level of the atlas. The external carotid artery gives off the following vessels. The a. occipitals (88/2), ranning towards the head, supplies the dorsal neck and nape musculature with a superficial and a deep ramus, and anastomoses with branches of the a. vertebralis and the a. comes vugi. The «a, sublingualis (/6) takes a course past the larynx, enters the mandibular space and then proceeds towards the tip of the beak, giving off branches to the cranial larynx, the submaximillary and Lingual musculature, the salivary glands and the lower beak. The a. laryngica propria which carries blood to the larynx and trachea and the a. oesophagica descendens (17) cattying it to the osso- phagus and trachea, arise as a common trunk from the sublingual artery. ‘The a. byoidea (/8) originates from the external carotid at the same level as the sublingual artery. Ascending towards she nape region it crosses the a. carotisint., follows the large ramus of the hyoid bone and supplies the musculature of the hyoid bone, the lower jaw and of the nape region, as well as the skin (of this area. Medial to the mandibular joint the a. lingual leaves the external carotid artery to ramify in the tongue. The a. mandibularis(/9), a. palatina (/10) and a. pterygoidea arise near the origin of the lingual artery: The mandibular artery proceeds along and below the ramus of the mandible and, with one of its subsidiaries, the a. alveolaris mandibulae, conveys blood to the muscles of the mandibular space, the salivary glands and the mucous membrane of the mouth cavity; in che fow it also supplies the wactles. The stout palatine artery is initially situated between she masticatory muscles whence its course is directed below the mucous membrane of the palate to the point of the beak. Apical to che choanal cleft the two arteries from each side unite into 4 single vessel which enters the beak and continues to the point giving off branches along its course. The slender pterygoid artery ramifies in the median masticatory muscles. The a. auricularis (21) springs from the external carotid as the latter lies alongside the quadrate bone and it supplies vessels to the m. depressor mandibulae, the external auditory meatus and the surrounding region. ‘The a facialis(/12) represents the continuation of the external carotid artery. Ie continues past che quadrate bone and through the depressor muscle of the mandible, progressing subcutaneously and parallel to the zygomatic bone in the direction of the upper beak. Ie then rises co the nasal border of the orbit to reach the forehead where, in the fowl, it fans out vo supply the comb. Along its course it provides several branches to, among other structures, the depressor muscle of the mandible and the eyelids. ‘The a. carots int. (88/5) ascends to the base of the skull, perforates the occipital bone ventra- laterally to the for. occipitale magrum, and divides into the a. opbthalmica (/13) and the a. carats cerebralis (|14). The ophthalmic artery circles round the caudal contour of the external auditory meatus and rakes 2 course towards the orbit. From it spring the slim a. temporalis, running to the94 Aneriel Fig tH, Arteriey of the head of che foel Ase Grsias 939) 5 2 ceophagla docs @'t hyoides; 2. mandbelaris oa pinay 8, ‘acl 72, 72 a. fal, ss ophbaimias 10a coc cee muscles of the same name and co the skin of that region, and the a. alveolaris inf. which enters the mandibular canal in company with the mandibular nerve. The ophthalmic artery supplies small subsidiaries to the eyelids and, together with the «. ophrbalmica int., a branch of the cerebral carotid artery, it forms a vascular ring around the optic nerve from’ which arterial blood is conveyed t0 the bulbus oculi, the eye muscles and the lacrimal gland. Traversing the carotid Canal of the sphenoid bone, the a. earotis cerebralis eaches the base of the brain in the cranial cavity. Lateral to the pituitary gland it divides into the 2. ophthalmica int., which enters che orbit swith the optic nerve, and into a ramus rost. et cand. The ramus rost. gives off the a. cerebri prof. ‘which dips down between the midbrain and the rhombencephalon, ‘The ramus then forks into the «. cerebralis nasalis and the a. ethmoidea. The former makes a laterally convex curve round the base of the cerebrum which it supplies with numerous subsidiary vessels. The ethmoid artery reaches the orbit where it anastomoses with the external ophthalmic artery. Leaving the orbit avits nasal border, it enters the nasal cavity where it vascularises the nasal and rurbinate mucous membranes. Caudal co the hypophysis, the rami caudd. of the a. carotis cerebralis unite to become thew. basilais. The latter lies along the base of the shombencephalon and, apart from giving off several smaller arteries to either side, it gives rise to the a. cevebelli cand. This ascends the lateral surface of the cerebellum and also conveys the blood by small vessels to the inner ear. Tt continues over the medulla oblongata and finally terminates as the . spinalis ventr ‘The a. subclavia (96/10; 87/7), as already pointed out, takes its origin from the brachiocephalic artery. Its course is in a craniodorsally directed curve to reach the costal process of the sternom ‘and provide the arteries of the pectoral region and the wing. Its first subsidiary is the «. sterno- clavieularis (86/13; 87/10), which crosses the coracoid bone to the thoracic inlet where it divides nto the a. stermalis (87/11) and the a. clavicularis(/12) to the supracoracoid and pectoral muscles. ‘Another offshoot from the subclavian artery is the a. acromialis (86/14; 87/13) which proceeds to the shoulder joint between the coracoid bone and the coracobrachial muscle. It supplies not only the shoulder joint but also the coracobrachial muscle and the caput longum: of the triceps brachii ‘After giving off the axillary artery, the subclavian arcery continues as the short a. thoracica from ‘which arises the a, thoraciea int. with its ramus dors. e¢ ventr. (86/15, 16). Both these rami run to the inner surface of the thoracic wall, the dorsal branch following the ventral border of the lung while the ventral continues at the level of the sternocostal joints. ‘The next vessel is the «4. thoracica ext., from which spring the a. thoracica ext. dors. et ventr. (86)17, 18; 87/14) going to the pectoral muscles. Another branch of the thoracic artery is the a. thoracica lat. (86/19; 87/13) which reaches the dorsal border of the supesficial pectoral muscle and supplies particularly those parts of the skin where the so-called incubation patches develop at the time of brooding. In birds the a. axillaris (86/20; 87/16) is a branch of the subclavian artery and not, as in mammals, the continuation of that vessel. Tes first thin offshoots are the a. subscapularis and the 4. coracoidalis. The former supplies the subscapular and the ventral serratus muscles and the latter the shoulder joint and the coracobrachial and teres minor muscles. From then onwards the axillary artery continues as the a, brachialis (86/21; 87/17), passing over the caput longum of the ‘mn. triceps brachié into the groove between the latver muscle and the m. biceps brachii and dividing at about the middle third of the upper arm into the a. ulnaris (86/225 87/18) and the a. radialisted of the fowl wae felis She centre inf. which enters ic artery supplies ach of the cerebral arterial blood is axsing the carotid ain in the cranial ch enters the oxbie he a cerebri proj. vs then Forks into vex curve round he ethmoid artery ing the orbit ac its turbinare mucous is unite to become i from giving off ascends the lateral x ear. It continues he brachiocephalic sof the sternum y is the a. sterno- f where it divides 1 pectoral muscles. ‘which proceeds t0 supplies not only the triceps brachii 1. thoracica from oth these rami run tral border of the next vessel is the 8; 87/14) going to lat, (86/19; 87/15) particularly those seding. ry and not, as in scapularis and the s muscles and the then onwards the ‘put longum of the ‘ach and dividing and the a. radials Aorts"deseendens 95 (86/23; 87/19), Prior wo this division however, it gives off the a. profunda brachii (86/24; 87/20) ar the level of che shoulder joint. This vessel courses between the heads of the triceps muscle to supply it and the metapatagium of the wing. After the a. humeralis (87/21) leaves the deep brachial artery to supply the triceps and deltoid muscles, the latter vessel divides into the a, collaterals radials (86/25; 87/22) and the a. collaterals ulnaris(/26; /23). The former continues aiong the humerus beneath the triceps brachii muscle and then crosses the flexor aspect of the elbow joint to supply the musculacure and skin of that region. The larger a. collateralis ulnaris accompanies the m. triceps brachii dstad to traverse the extensot aspect of the elbow joint. Apart from supplying the muscles of the upper arm, it carries the arterial supply to the primary feathers and che skin of this area. A further branch of the brachial artery, the a. circumflex bumeris ant (27; 124), enters the biceps brachii muscle at about the proximal third of the humerus and supplies it and the propatagium. Along its course towards the elbow joint, beeween the biceps and triceps muscles, the ulnar artery is accompanied by the corresponding veins and the median nerve. It traverses the flexor aspect of the elbow joint and continues on the flexor carpi lnaris tusele to the carpal joint. The a. recurrens ulnaris (87/25), which is a subsidiary of the ulnar artery, crosses the medial surface of the elbow joint; it supplies blood to this join, to the flexor carpi ulnaris muscle and to the primary feathers of this region. Tt finally anastomoses with the 4, collaterals whnars. In the region of the carpal joint the a. wlnaris splits into several smaller vvessels which are distributed to the metapodium and acropodium. The first course of the radial arcery is parallel with the a. wlnaris as far as the elbow joint where it crosses the terminal tendon ‘of the biceps brachii muscle and continues on che medial surface of the lower arm, between the flexor and extensor muscles, to divide into small twigs at the carpal joint. On the flexor aspect of the elbow joint it sends a stout branch to the extensor muscle of the radial carpal, the pronator ‘muscles and the propatagium. Aorta descendens and its branches ‘The aorta descendens (86/4', 4”; 96/1, 1") gives rise vo paired aa. intercostales which issue at regular intervals from the 5th intercostal space onwards. Ascending dorsad, these intercostal vessels reach the appropriate intercostal spaces where dorsal brandies are given off to the musculature of the trunk and the vertebrae. Smaller offshoots enter the vertebral canal to contri- bute to the formation of the a. spinalis ventr. A ventrally directed branch divides into two vessels above the tuberculum costae; these descend on the eranial and caudal borders of the respective ribs to supply the external and internal intercostal muscles and ultimately to anastomore with branches of the internal thoracic artery. The aa. lumbales and the aa. sacrales respectively substicute for the intercostal arteries in the region of the synsacrum. ‘These arteries have also derived from bilateral, segmental arteries originating from the descending aorta in this region. Similarly, their dorsal limbs supply the trunk ‘musculature, the vertebrae and, by entering the vertebral canal, the spinal cord. ‘Their ventral branches vascularise the abdominal muscles and the skin of this area. ‘The aa. coccygicae are paired branches which arise from the a. coccygica media (86/28). The latter stems from the medial sacral artery. They have dorsally dizected subsidiary branches which convey blood to the dorsal musculature of the tail, the follicles of the “steering feathers”, the skin and the preen gland ‘The first large ventral limb of the descending or, as it may now be termed, the abdominal aorta is che a. coeliaca (86/29; 89/1). Immediately following its origin it gives tise to a ramus o€s0~ phagicus. The first main trunk of the coeliac artery is the a. gastrica sup. (86/31; 89/2). A ramus dors. (89/3) from the later supplies the right and the dorsal surfaces of the proventriculus, a ramus hepaticus (/4) supplies the left lobe of the liver while a ramus intermedius (/5) goes to the left wall of the gizzard. The second main trunk of the coeliac artery is the a. gastrica in). (86/32; 89/6) which provides several small vegels to the spleen (89/7) and, at the same level, a ramus bepaticus dext.(/8) to the liver. The a. gastrica inf i continued as the a. gastropancreaticoduode- nalis (/9) from which springs 2 branch going to the ascending limb of the duodenum (/10), a ramus ‘gastrieus dest. (/11) to the muscular stomach, and rami ileocaecales (/12) to the middle section of the caeca and the ileum situated between them. The gastropancreaticoduodenal artery then con- tinues as the «. pancreaticoduodenalis (/13) which pursues a course along the pancreas between the ‘two limbs of the duodenum, which organs it supplies by means of several subsidiaries.9% Arveries Buganede epesenston fo pe 1 beni of theaters « el porns 3,24 le ene dere. a i Tt hdindes toe dene and Gon Berl 3 a reali ora det 4 onion emf Se a saa eas SI cen dest tod ‘The a. meventerica eran (86/30; 89/14) springs from the Stedominal aorta near the or- in of the coding arcery. The Fite: vee arising. from the sieocaeels (89/19) which Sasculriss the fis part of the Geet andthe ileum. ‘The cranial mesenteric artery con- (16), from whith numerous vessel, the ram eats (17), trseatshortinverval Thelatet convey Blood 10 the jejunum The a. mesentericaeaud, (86/33; 8911; 91/19) i a relatively small vessel which does nr leave the abdominal aorta unt the fst pat ofthe pelvic cavity atthe level ofthe caudal renal lobes Te vides into evo limbs (99/1, 20) one of which, after giving off numerous brandet 0 the Colon fellows a cranial course inthe meocolon to anastomore with thes eoeatali. The cond Timi fans out on the colon Ac the level ofthe ltt choracc or first lumbar vertebra, the abdominal aorta of the co gives off wo each side a common sem, comprising both the 3 renais eran. and che a. spermatia {hts testculars (91/2). The later sends several wel o che tests while the canal renal artery supplies the cranial lobe of the Kidney with numerous subidary arere; some small branche also go to the adrenal gland (86/34; 91/9). la female Bids there is'no veel on the right side corresponding 1 the spermatic arvery, s0 this vel is own only se the a. reals cra, dest (9015) However on the left side in female bird very thid stom originate fom the aorta, and this divides ino an a, spermatica int. ovaiea(26/94"; 9018") an. venlis sin cram (86/945 Bods") and an a. ovidnea eran. (86136"", 9019"), The a ovarce brand in the’ fheovaiu,she anirs¢ cok en dere und 1114) springs from the L aorta near the ori= coeliac artery. The el arising from the csenteric artery is che lis (89/15) which ce the firse part of the d the ileum. ‘The ceenteric artery con the truncas jejunalis vm which numerous fe rami jejunalis (/17), ortintervals. Thelateer food to the jejunum. which does not leave he caudal renal lobes. erous branches to the ceacaecalis. The second aorta of the cockerel and the a. spermatica se cranial renal arcery some small branches el on the right side 4. renalis eran. dext. ss from the aorta, and ualis sin. cran, (861345 cs in the mesovarium, PLATE IV (afer Voruartsaun apebied wok anode athe lana of Vausaary Aaseony, Gen) A cri vreens B epg verhen; © pity D endeay E beat; Phas @ hier eveys J ee dee K thyroid gland Z ccehalica in. and’ dene @ a sci media: 9 a, ec tas stot f2 cmos wal areal tara olny 24 a pro. ra 35 ‘isan end vw. cnaphaseee ‘eis sferes dud in fires dtd ss 69 v= excygmameseias 6PLATE VPLATE VIPLATE VIIod locas Gant discon gm wenieenbrne "oP nous mucin 2 sta afree dee ted int Teed, (2); 3m peroment, Arseries of the hind limb 7 thus reaching the ovary from dorsal. The 2. oviducts curves caudally, enters the suspensory liga- ‘ment of the oviduct and supplies the infundibulum and cranial pare of the isthmus. The ¢. renalis sin. is responsible for the arterial supply to the left cranial lobe of the kidney. Arteries of the hind limb ‘The hind limb of the bied is supplied with blood by che a. iliaca ext. (86/33; 90/2; 91/6) and by the a. ischiadica ext, (86/36; 90/3; 91/7) ‘The a. iliaca ext. arises from the aorta on the ventral surface of the synsacrum between the two riddle lobes of the kidneys and it leaves the pelvie cavity above the hip joint. Ie first branch is the a. circumflexa femoris (86/37; 91/8) whic arises at the level of the bip joins. From this circumflex vessel springs the «. glutaes (/38; /9) which proceeds towards the outer surface of the ilium and the gluteal muscles, thence distally to the cranial surfaces of the thigh to supply the extensors of the knee joint. The main stem of the external ise artery crosses the lateral surface of the hip joint where it divides into the a. pelvina (/89; /10) and the a. femoralis (86/40). The 4, pelvina follows the pubic bone caudoventrally giving off vessels to the abdominal air sac and the musculature of the pelvis and abdomen. In females the a. pelvina also gives rise to the 4, oviducts media (86/39). On the medial sarface of the thigh the a. femoral rons down to the flexor aspect of the knee where it becomes the 4. genus suprema. Both these arteries supply the knee joint and the muscles of the medial aspect ofthe thigh, especially the adductor. ‘The aa, ischiadicae extt. (86/36; 90/3; 91/7) take their origin from the aorta at the level of the hip joint. They traverse the left and right kidney respectively in the groove between the middle and caudal lobes and leave the pelvic cavity through the foramen ischiadicums in company with the m ichiadicus. While still within the pelvic cavity, che a. ixhiadica sin. gives rise to the a. renalis media sin. (90/6; 91/11) and che a. renalis cand. sin. ((7; 112), either independently or from a common trunk, These vessels provide the arterial supply to the middle and caudal lobes of the Kidney respectively. In the majority of female birds che a. renalis media sin. gives rise to the 4 oviducta media (90/6") which lies in the suspensory ligament of the oviduct. Tt supplies the middle part of that organ and then anastomoses with branches of the 4. oviducta eran. In a small percentage of cases it issues directly from the aorta together with the left caudal renal artery. The middle and caudal lobes of the right kidney are supplied by the a. renalis media dext. and the 4. renalis caud. dext. respectively; these derive in a common trunk from the right ishiatic artery. After traversing the ichiatic foramen, the ischiatic artery gives off muscular branches and takes a course, together with the ischiatic nerve, caudal to the femur between the criceps femoris muscle fon the one hand and the semimembranosus and adductor muscles on the other, tothe flexor aspect of the knee joint where it becomes the a. poplites (86/313 91/13). During this course the «.trochar= terica (86142) is given off and runs cranially to the hip joint. Another artery follows a caudal direction to supply the obturator, quadratus femoris, biceps femoris and gemellas muscles. Another branch, the a. profunda femoris (86/43; 91/14) supplies the biceps femoris, che semicendinosus, the semimembranosus and the adductor muscles. A main trunk, the subsidiaries of which anastomose with similar arteries from the a. profonda femoris and the a. tibialis med, leaves the popliteal artery and thereafter gives rise to a. femoris caud. (86/44; 91/13). Branches from the latter vascularise the flexor muscles in the lower leg and the skin of this region. The a. tibialis med. (86145; 91/25) also stems from the popliteal artery. Leaving the parent vessel on the flexor aspect Of the knee, it takes a medial course down the lower leg under the gastrocnemius muscle then, making 2 dorsally convex curve, it continues distad to participate in the formation of the rete tarsometatarsicns Ie cartes blood to the muscles on the dorsolateral aspect of the lower leg. Also originating from the popliteal artery is the a. tibialis post. (86/46). Tt follows the caudal aspect of the tibia in a distal direction between the m. flexor hallucis longus and the m. flexor digitalis pedis prof. and also joins the rete tarsometatarsicus. Arising from the popliteal artery at the same level as the posterior tibial artery, is the a. peroneal (91/28) which rises to the dorsal surface of the lower leg where ic supplies the flexor muscles of the tarsometatarsal joint and the extensors of the toe joints. Having given off all these branches, che a. poplites finally divides in the middle third of the tibia into the stout a. cialis ant. (86/47; 91/17; 92/1) and the smaller a. tibialis lat. (86/48). The latter descends distad in a lateroplantar position in the groove between the tibia and fibula, is deflected onto che medial side of the art. tarsometatarsica, to supply that joint and