APPENDIX DEVELOPMENT OF CHICK EMBRYOS TO 4 VOS FROM 18 HOURS YS OF INCUBATION STUDY OF SERIAL SECTIONS To study embryos with any degree of thoroughness one must cut them into sections which are sufficiently thin to allow effective use of the microscope to ascertain cellular organization and detailed structural relationships. In the prepa- ration of such material the entire embryo is cut into sections which are mounted on. slides in the order in which they were cut. A sectional view of any region of the embryo is then available for study. In studying a section from a series, it is first necessary to determine the location in which it was cut through the embryo. The plane of the section under consideration, as well as the region of the embryo through which it passes, should be ascertained by comparing it with an entire embryo of the same age as that from which the section was cut. Only when ie location of a section is known precisely can the Rie ts ie correlated with the organization of the embryo as a who aaa iaate study of embryology causes students more difficulty than aoe ae te sections accurately, with the consequent failure to appreciate the relaiovens © the snacturee seeg in them, The importance of fiting the structures shown by sections properly into the general scheme of organization as it appears mounts (Figs. A-l and A-2) cannot be overemphasized. TERMS OF LOCATION In embryology it is necessal direction of growth processes Oy ies. embryo regardless of the position it occupies. tefer to the direction of the action of gravity, ry to designate the location of stein and the i ferable to the body of the by terms rc ur ordinary txms of location rr, and under. 629 swuch as above, Ove Dipindai dengan CamScanner5 OF INCUBATION ~ 630 pevexopMenr oF cHiex eManvOs FROM 18 HOURS TO + OY Plane of section LONGITUDINAL SECTION. ‘TRANSPARENT PREPARATION E ‘OF ENTIRE HEAD FIGURE At | Relation of longitudinal section of the embryonic head to the picture presented by a head of the same age mounted entire as a transparent preparation. ‘These terms are not sufficiently accurate in this type of work because the embryo itself may lie in'a great variety of positions. The correct adjectives of position are dorsal, pertaining to the back; ventral, the belly; cephalic, the head; caudal, the tail; mesial, the middle part; and lateral, the side of the embryonic body. In dealing with relations to the head, the adjective cephalic is inadequate and the term rostral (Latin, beak of a bird, bow of a ship) is used to designate the extreme anterior portion of the head or the relative location of intracephalic structures such as the various parts of the brain. Adverbs of fixed position are made in the usual way by adding -ly to the root of the adjective. _In addition to adverbs of position, corresponding adverbs of motion or direction are formed by adding the suffix -ad to the root of the adjective, as in dorsad, meaning toward the back; cephalad, meaning toward the head, etc. These adverbs should be applied only to the progress of processes or to the a ie einetites eee the. part indicated by their root. Thus, for ese fe fae iat the developing eye of an embryo was located in the aie Near ae aon or that the forebrain was in the cephalic part of the it moves farther tortor eaie: CxPress the idea that as the eye increases in size » We would say it grew laterad. Fro THE PRIMITIVE-STREAK STAGE TO THE APPEARANCE ee Stages of embryonic development ofthe chick occur within ‘act of the .hen, the laboratory study of chick embryos Dipindai dengan CamScannerFROM THE PRIMTIVESSTREAK STAGE TOTHE APPEARANCE OF SOMES 631 Inthe entire mount these lines ‘appear most clearly au folds, high focus Margin of gut, ride depth Edge of mesoderm, low focus ‘TRANSVERSE SECTION FIGURE A-2 Flin of ansverse section of the embryonic head to the picture presented by an fete head of the same ago viewed as a transparent preparation. Commonly omits the early stages of cleavage and blastulation, Gastrulation and the formation of the primitive streak were described in Chap. 5 (Figs. 5-12 to $16) and will not be repeated here. By 18 hours" incubation the primitive streak 'S well established, and cells which have been invaginated by way of Hensen’s node push cephalad, initiating the formation of the notochord (Fig. 5-18B). In hick embryos which have been incubated about 18 hours the notochord has ecome markedly elongated to form a well-defined midline structare (Fig. A-3). welder embryological treatises the young notochord was frequently called the cen sees and chick embryos of about 18 hours’ incubation have often poken of as being in the “‘head-process stage.”” Sections of embryos in the aac stage (Fig. A-4) give much more ‘nformatioy hord than can be sleaned from tet the structure and relations of the notoc cil 's node, the the study of whol its. Near its origin at Hensen's m Totochora y ly of whole mounts. ally with the Tce is made up of diffusely arranged cells that merge cau qos vand the 'urned under cells of the lateral portions of the mesot Dipindai dengan CamScannerS82 DEVELOPMENT OF chick EMBRYOS FROM 16 HOURS TO 4 DAYS OF NCUBATION Ceohalie end Prosmnion Amerior border of mesoderm Neural pate — Notochord Emboryona are. enters node Btrieve WA 5S primitive nresk Area pellucida Area opace five Oe FIGURE 43 . Dorsal view (14) of entire chick embryo of 18 hours’ incubation. 7 Caudal end endoderm. Its cephalic tip graduall rodlike configuration, In the sections diagrammed in Fig. A-4 a conventional scheme of represen: tation has been employed to indicate each of the germ layers. The ectoderm is vertically hatched, the cells of the mesoderm are represented by heavy angular dots when they are isolated or by solid black lines when they lie arranged in the form of compact layers, and the endoderm is represented by stippling backed by a single line. A similar conventional representation of the different germ layers is observed in all diagrams of chick sections to facilitate following the way in which the organ systems of the embryo are constructed from the germ layers. The plane in which each of the sections diagrammed passes through the embryo is indicated by a line drawn on a small outline sketch of an embryo at ‘@ corresponding stage. ly develops a more definitely circumscribed, The Primitive Streak Dipindai dengan CamScannerEctoderm of Primitive gut Primitive Ecraderm pes Som anton age te? Endoder “Area of o indiferent eos "GH POWER THROUGH PRIMITIVE STREAK, HIGH POWER THROUGH EDGE OF AT REGION 0 ON SECTION C, GERM WALL AT REGION E ON SECTION C. Heren’s node rnnn —_ewtpne | iin ee, eine 90 1 Notocnord Endoderm tens : h Extent of primitive gat and of 0a pace Ea “ 1 etic grat tick The location of each section is incated by ale drawn cn a small ut eae ti eee embryo of eonesponcing ‘age. The letters affixed to the lines on the sketch ean ayer ig gest Sections correspond with tho letters designating the socton diagrams. tenied by a dierent conventional scheme: ectoderm by verical hatching; by a single line; the cells of the mesoderm, which at this stage do Yee chord fy op NeSYY angular dots, (A) Diagram of transverse secon trough neural (522 Brough preadt™ Of transverse section through primitive pit. (C) Diagram of rans- SRW showing Ye SHAK. (0) Draving showing collar suc in pritivesreak e- Section Cellular structure at inner margin of germ wal (F) Dlagram of median Passing trough notochord and primitive steak. Dipindai dengan CamScanneri 834 DEVELOPMENT OF CHICK EMBRYOS FROM 1S HOURS TO 4 DAYS OF INCUBATION length of strerk oo Post. border of area palocice Hours 0.0 FIGURE A-5 of the notochord. The at the left. The time is, the ever-elongating notochor al half of the streak contributes to the lateral and caudal expansion of both the intra- and extraembryonic tissues and also accounts for much of the shortening of the streak. As the primitive streak d (Fig. A-5). Emigra- Dipindai dengan CamScanner635 FROM TH PRIMNTIVE-STREAK STAGE TO THE APPEARANCE OF SOMITES Cute for emevsl of enbrya = Yolk = no verre Messcape sy macs we ante 2 epee ea SS neta cone ga Sa bodies spread out on a large mass of yotk, a aaa daainel a is known as the germ wall (Fig, A-4E). The cavit endoderm is now termed the primitive gut. Whe sectioning, the yolk floor of the primitive gut, not being adherent to the blastoderm, is left behind (Fig. A-6A). In contrast, the periphery of the blastoderm lies closely applied to the yolk. Some yolk adheres to the part of the blastoderm when it is removed (Fig. A-6A). Its presence clearly indicates why the area opaca appears less translucent than the area pellucida in surface views of entire embryos (Fig. A-3). One can obtain a better sense of perspective on the relations of the flat avian embryo by anne it would appear if the embryo were folded ventrally into a cylinder (Fig. A-6). . By about the twentieth hour of ination we ca st indeatons of ea diferentaton of the region ofthe primitive gt ha under he cote pat of the embryo. The first part of the gut to acqui . Pocket is called the foregut (Fig. A-7). Growth and Early Differentiation of a men the primitive streak spreads i ise§ from either si z oderm swings pees Sarai the same time each lateral wing of the mes‘ Dipindai dengan CamScanner'S OF INCUBATION 636 _Devevopwten oF CHICK EwBAYOS FROM 18 HOURS TO 4 DAY Eetoderm of herd ie end Prosmnion « Coonat aig, Borde of foregut Subeeohatc pocket . ‘ TSS Neural groove 3 Hensers noe Ccoudlend 4 FIGURE 4.7 Dorsal view (X14) of entre chick embryo of anout 20 hours’ incubation, cephalad (Fig. A-8A to A-8C). The manner in which the mesoderm spreads out leaves a mesoderm-free arca in the cephalic portion of the blastoderm. This region, known as the proamnion, is clearly recognizable in entire embryos by reason of its lesser density (Figs. A-7 and A-9). Despite the implications of its ‘name, this region is not the precursor of the amnion. Dipindai dengan CamScannerPrearnion Notochord = staaan's node A CHICK o| F ABOUT 14 Ho URS 8 chick OF ABouT 18 HOURS Anterior horn of mesoderm Proamaion Primitive streak E CHICK OF ABOUT 16 HOURS THE SAME INJURED AT THE cHick THREE POINTS MARKED AFTER FURTHER INCUBATION FIGURE A-8 itive streak (AC) show the progress of the Diagrams outing direction of growth trom th pri dem ‘during the latter part of the first day of incubation. The areas into which the meso- dorm has grown are incicated by diagonal hatching. (D) oo te which gives rise to the spinal cord. Three primary brain vesicl 7 ce distinguished in the enlarged cephalic region of the neural tube a from it bY most rostral vesicle is the forebrain, or prosencephalon. Marke ol mrt a constriction is the midbrain, or mesencephalon. A very slight constr marks the boundary between the mesecephalon and the Hinde o oe encephalon, which is continuous with the future spinal cont) the late definite point of transition. In somewhat older embryos (Fig. Dipindai dengan CamScannerCHANGES BETWEEN 24 AND 23 HOURS OF WouBATION 645 Proamnion —_Prosencephalon Border of foregut Anterior eal ay Mevenceahalon ‘Subceohalic pocket Notochors Metenchy ne Vitetine Rhombenesphalon Margin of antatior Intestin! poral Somite Sinus homboitie. + Hensens noge yokes yee Primitive area ee vetcular plows ricune ats 3 Dorsal view (x14) of the entire chick embryo having 8 pairs of somites (about 27 to 28 hours’ incubation). walls of the prosencephalon become outpocketed to form a pair of rounded dilations known as the primary optic vesicles. Ventral to the brain the notochord extends rostrally as far as a depression in the floor of the prosencephalon known as the infundibulum (Fig. A-16). As the result of an interaction with the ectoderm of the future oral cavity, the infundibulum develops into the neural hypophysis. ‘The closure of the neural folds takes place first near the rostral end of the neural groove and then progresses both cephalad and caudad. Closure of the extreme rostral end of the brain is delayed, and the persisting communication between the prosencephalon and the outside is called the anterior neuropore (Figs. A-15 and A-16). By 33 hours it is almost closed (Fig. A-18), and later it becomes entirely closed, leaving only a scarlike fissure in the anterior wall of the prosencephalon, At the other end of the body, the neural tube has closed caudally as far as somite formation has progressed. Caudal to the most posterior somites, the neural groove is still open and the neural folds diverge to either side of Hensen’s node (Fig. A-15). The lateral boundaries of the unclosed region at the posterior end of the neural tube delimit an open area known as the sinus Dipindai dengan CamScanner546 DEVELOPMENT OF CHICK EMBRYOS FROM 18 HOURS TO 4 DAYS OF INCUBATION [Anterior horn of mesoderm Proamnion: VR Eetoserm of hess "Amir neronre prueznoon re Optic venice aE Cephalic mesenchyme Notechord Ventral aortic root Mezencephslon Ventral sort Epimyocardiom Margin of foregut Enéocarsium Rhombenceohalon Line of encoesral t Viteltine vein thargin of aterior { ‘metnal poral <7 Enrmembryon vse pens Somixe FIGURE A16 \Ventral view (x47) of cephalic and cardiac region of chick embryo of 9 somites (shout 29 to 30 hours" incubation), rhomboidalis (Fig. A-18). Hensen's node and the primitive pit lie in the floor of this still unclosed region of the neural groove; subsequently they are enclosed within it when the neural folds here finally fuse to complete the neural tube. ‘This process in the chick is homologous with the enclosure of the blastopore by the neural folds in lower vertebrates (Fig. 5-4E). In forms where the blastopore does not become closed until after it is surrounded by the neural folds, it for a time constitutes an opening from the neural canal into the primitive gut known as the neurenteric canal. In the chick the fact that there is never an open blastopore precludes the establishment of an open neurenteric canal, but the primitive pit represents its homologue. ‘The Foregut and Heart The anterior intestinal portal occupies progressively more caudal positions (Fig. A-22) as the margins on either side of the portal are constantly converging toward the midline. Such merging, along with elongation of the structures cephalic {to the anterior intestinal portal, results in lengthening of the foregut. The resulting enlargement of coelomic space between the subcephalic pocket and the margin of the anterior intestinal portal is accupied by the developing heart. Dipindai dengan CamScannerSTRUCTURE OF CHICKS BETWEEN 29 AND 38 HOURS OF NCUBKTION 647 ce, a dorsal View Phalon 5 Fig. Paited viteline van VeSS2h the ventral aorta STRUCTURE oO} INCUBATION F CHICKS BETWEEN 33 aND a9 HOURS OF The Brain and Its Neuromeric Structure The metameric arranjgement of structures, which is 50 striking a feature in the body organization of all vertebrates, is masked in the head region of the adult by superimposed specializations. In the brain of young vertebrate embryos, however, the metamerism is still indicated. Dissections of the neural plate of chicks at the end of the first day of incubation show a series of It enlargements marked off from each other by constrictions (Fig. A-17). With the closure of the neural tube and the establishment of the three Primary brain vesicles, we can begin to trace the fate of the various neuromerie enlargements in the formation of the brain regions. The yee a vated Reuromeres form the prosencephalon, neuromeres IV an are ear a into the mesencephalon, and neuromeres VI to XI rae interneuromerie thombencephaion (Fig. A-17B). Rostrally, all but oo ra Constrictions soon disappear, namely, the one between hada hd the rhobea- the Mesencephalon and the one between the mesencepl remain clearly marked Cephalon. The rhombencephalic neuromeres, however; for a considerable period. Dipindai dengan CamScannerG48. DEVELOPMENT OF CHICK EMORYOS FROM 18 HOURS TO 4 DAYS OF INCUBATION — ior neurepore Nera groove sn Lett near fols Somite Proseneephalon Jao \ f-sotn iyeleneephaion postion of auditory pit ch emo (Os Denar 25 0 27-hour) embryo. (C) Dore vena pa we ol rain is mye. (0) Dorsal view of raln tom 1¢somte (Soha) ‘embryo. (Based on figures of Hil) By about ir ; a eae eer 7 incubation the optic vesicles are established as paired Width ofthe head Figs reseteePhalon, They soon extend to geeupy te full 8s. A-ITC and A-18). The distal portion of ench of the Dipindai dengan CamScanner!RUCTIRE OF C¥EAS BETWEEN 3944038 HOURS OF neagn ion 649 Hart sumencephton Optic vetice Border of tovegut Vittiog vin Rrombenesahaton Notoenora Neural tube Sinus terminalis Sirus namboians Pete sea Figure are/ 5 “sal view (17) of an ent chick embryo of 12 somites (about 39 hours incubation. reales thus comes to lie closely approximated to the superficial ectoderm, a "lationship of importance in the later development of the lens. At sy ie “wities of the optic vesicles are broadly confluent with the cavity of the Dipindai dengan CamScannerPresmnion Prosenerphalon Ectederm te vette Uondioati oa omic ach ental sone Notochord Chale neural erat region of gangion V arg of forege Mereneephalon Truneus anesions yeteneephalon : é versie Cephate neural rest, SE ‘pion of goation| ee — susiory ot Vina vn ee: ae tyeteneenton : FIGURE 4-19 arsa! view (*S0) of cehale and cardiac regions ofa chick embo withthe ‘seventeenth so- ‘ite just forming (zbout 38 hours incubations” prosencephalon. Somewhat later, constrictions mark more definitely the boundaries between the optic vesicles and the prosencephalon (Figs. AcI7D and A-19). The infundibulum remains os a depression in the floor of the Prosencephalon (Figs. A-19 and A-20), In chicks of about 38 hours, indications of the impending division of the three primary vesicles to fc are already beginning to Dipindai dengan CamScannerTon 654 STRUCTURE OF CHICKS BETWEEN 39 AND 28 HOURS OF INCUBATION Anterior neureore _———— Prosercephaton Optic vesicte <— Infundibulum ut eoimyocareiom = ) Tomer | — Vento repion : Donal sons Text Anterior internal porta! Vitetine vein Cut splanchnopieure ‘Lateral mesoderm Notochord ‘nts Dorsal aorta Neural ube pebhalon. By 38 hours of incubation (Fig. A-19) the auditory placodes have ‘Come depressed bel; iad ed below the general level of the ectoderm and form the walls aude of Cavities, the auditory pits. When first formed, the walls of the cavitin” Pits are directly continuous with the superficial ectoderm and their les are wide Open to the outside. In later Stages the openings into the pits NE Rarrowed and finally closed, so that ine pits become vesicles lying Dipindai dengan CamScanner652 _ DEVELOPMENT OF CHICK EMBRYOS FROM 18 HOURS TO 4 DAYS OF INCUBATION — between the superficial ectoderm and the myelencephalon. At this point they have no connection with the central n lervous system. Formation of the Heart The heart arises from paired primordia, which at first lie widely separated on cither side of the midline. The paired condition of the heart at the time of ite origin is due to the fact that the early embryo lies open ventrally, spread out on the yolk surface. The primordia of all ventral structures which appear at +n carly age are thus at frst separated and lie on either side of the midline. As the embryo develops, a series of foldings undercut it and separate it from the yolk. At the same time, this folding-off process establishes the ventral wall of the gut ang the ventral body wall of the embryo by bringing together in the midline the structures that were formerly spread out to right and left. As the embryo is completed ventrally the paired primordia of the heart are brought together in the midline and become fused (Figs. A-21 and A-22). Although prospective cardiac tissue can be recognized much earlier, the frst definite structural indications of heart formation appear in chicks off 25 to 26 hours in the region of the anterior intestinal portal. Where the splanchnopleure of either side bends toward the midline along the lateral margin of the intestinal Portal, there is a marked regional thickening in the splanchnic mesoderm (Fig. 4-214). This pair of thickenings indicates where there has been rapid cell proliferation preliminary to the differentiation of the heart. Loosely associated cells can already be seen somewhat detached from the mesial face of the mesodermal layer. These cells soon become organized to form the endocardial primordia ~ Ina chick of about 26 hours, sections through a corresponding region show distinct differentiation of the endocardial and epimyocardial primordia (Fig A-21B). The endocardial primordia are a pair of delicate tubular structures, with walls only a single cell in thickness, lying between the endoderm and mesoderm. They arise from the cells migrating out of the adjacent thickened mesoderm in the 25-hour chick. As U their name indicates, they are destined to form the internal lining of the heart (endocardium). The greater part of each of the original mesodermal thickenings becomes applied to the lateral aspects of the endocardial tubes as the epimyocardial primordium, which will later differentiate into the external coat of the heurt (epicardium) and the heavy muscular layers of the heart (myocardium). Careful study of the developing heart at this stage reveals the presence of cardiac jelly between the epimyo- cardium and the endocardium, — In chicks of 27 hours the lateral margins of the anterior intestinal portal have been undergoing concrescence, thus lengthening the foregut caudally and at the Same time elongating the pericardial region. In this process the lateral margin’ of ae Derk ae a to meet each other and merge in the midline, and Le endocardial tubes of the right si i acl other beneath ee ight side and the left side are brought toward | foregut (Figs. A-21C and Dipindai dengan CamScannerNeural plate ectoderm “cephalic mesenchyme supe ectoderm Gator aus ee 5 So Fencaast 2) Somsteoiert * ‘eon of ) Splanehnpeure ‘fesse Notocnors Emerging eas Neal oor cephalic mevnehvne Engoearal primorcivm of anterioeintetinat Somatoplevre Splanehnepleure pimyocardal primer YPericarcial orion, Focegut Line of fsion of margins Sf anterior intestinal poral NX Neural canal Endoesraa! primordim oo } Somatooleure Endocardial primordium Foregut carcise itv Dorsal mesocardium (Horeing) Epimyocarium })Somatopleure ye, Perea toelom } splanchnopleure ° enna mesoearaium a Foreaut Enaceaeprimorcium Neural ube Interline Cardi jet Doral nietoearium Epimyecartim x — — ] Somatopleure ZL Pacarsl ‘ . % cevlom i @ } Splanchnopleure FE ene nescence {diappeina) Ficune a2 ¥ ol transverse sections through the pericardial region of chicks at various stages to on formation of the heart, For location of the sections, consult Fig. A-22. (A) At 26 hours; ‘hours; (C) at 27 hours; (D) at 28 hours; (E) at 29 hours. + - Pert rk Ss Dipindai dengan CamScanner854 DEVELOPMENT OF CHICK EMBRYOS FROM 18 HOURS TO 4 DAYS OF INCUBATION Ton of cotom \ ‘Ventral aortic root Pericarsi\ region of eoelom Epimyocardium Vitelline vein FIGURE A-22 Ventral-view diagrams to chow the origin and subsequent fusion of the paired primordia ol the heart. The lines A, C, D. and E indicate the planes of the sections diagrammed in Fig. A-21A, C, D. and E, respectively. (A) Chick of 25 hours; (B) chick of 27 hours; (C) chick of 28 hours: (C) chick of 29 hours. A-22B). In the 28-hour chick embryo the endocardial primordia are approxi- mated to each other (Figs. A-21D and A-22C), and by 29 hours they fuse in thei midregion to form a single tube (Figs. A-21E and A-22D). At the same time the epimyocardial areas of the mesoderm are brought together, first ventrally (Fig. A-21D) and then dorsally to the endocardium (Fig. Dipindai dengan CamScannerwT ‘STRUCTURE OF CHICKS BETWEEN 39 ANO 38 HOURS OF INCUBATION 655 A-21E). Where the splanchnic mesodermal layers of the opposite sides of the body become apposed to each other dorsal and ventral to the heart, they form double-layered supporting membranes called, respectively, the dorsal meso- cardium and the ventral mesocardium. The ventral mesocardium is a transitory structure that disappears almost as soon as it is formed (Fig. A-21E). The dorsal mesocardium, although the greater part of it disappears in the next few hours of incubation, persists in embryos of the early part of the age range under Consideration, suspending the heart in the pericardial region of the coelom. The gross shape of the heart and its positional relations to other structures can be readily seen in entire embryos. The fusion of the’ paired cardiac primordia establishes the heart as a nearly straight tubular structure. It lies at the level of the hombencephalon approximately in the midline, ventral to the foregut (Fig. A-16). By 33 hours of incubation the midregion of the heart is considerably dilated and bent to the right (Fig. A-18). At 38 hours the heart is bent so far to the right that it extends beyond the lateral body ‘margin of the embryo (Fig. A-19). This bending process is correlated with the rupture of the dorsal mesocardium at the midregion of the heart. Although there are not yet any sharply bounded subdivisions of the heart, its fundamental regions are beginning to take shape. From the heart's future intake cad to its future discharging end, the regions are sinus venosus, atrium, ventricle, and truncus arteriosus. At the 36- and 38-hour stages the sinus venosus is only suggested. It is represented by the still paired primordia where the common cardinals enter the vitelline veins, and they in turn are becoming Confluent with each other to enter the atrial portion of the tubular heart (Fig. 4-25). The atrium is held close beneath the caudal part of the foregut by a Persisting portion of the dorsal mesocardium. The ventricle is the part of the cardiac tube that makes a U-shaped bend to the right, bringing it into clear view at the side of the body in whole mounts (Fig. A-19). Swinging back to the midline, the ventricle is narrowed to form the discharging part of the heart known as the truncus arteriosus (Figs. A-20 and A-25). From the way the paired cardiac primordia are at first located on either side of the anterior intestinal portal, it is evident that they can fuse with each other in a sequential process only as the “flooring in" of the foregut progresses (cf. Fig. A-21B to A-21D). The truncoventricular part of the heart is formed first (Fig. A-23A). Then the atrium is added caudal to the ventricle (Fig. A-23B and C). Finally, in stages older than those under discussion here, the sinus venosus is added caudal to the atrium (Fig. A-23D), Formation of Intraembryonic Blood Vessels it i ca vd vessels arise within the Concurrently with the establishment of the heart, bloos body of the embryo, The large vessels connecting with the heart are the first of the intraembryonic channels to be established. The endothelial lining of the truncus is continued cephalad beneath the foregut as the yentral aorta (Fig. A-25). Almost immediately the ventral aorta bifurcates to form the paired Dipindai dengan CamScanner1656 DEVELOPMENT OF CHICK EMBRYOS FROM 18 HOURS TO 4 DAYS OF INCUBATION Entrance ot common cardinal ‘cin marking torure sinus level venosut| ‘established o Ire md bidet The Vere! yw atthe Gearon’ OF the chick heart by progressive fusion of its paired cites pen oe esleQe (about 28-29 hours), when the fist contrac: orid the myocardet ayot'®, OF fegion where the fusion of tha paired ure) when ester has been formed. (B) Veriral view at the bat the enue ono fist begins to circulate, The atrium and hig mye noeUs &XIS only a5 undtterentiated primordia! Of Doe pavestment. (C) Verito-sinistal view at the 19° paired primordia is just beginning to involve the vith save (about 51-83 hours). The sinus venosus ayoearcium is well advanced. To taciitale alta at te po “en placed against approximately este cardinal von 95 3° (From Paten and Kram, 1850) oe Dipindai dengan CamScannerounces SETWEEN8 AND 50 HOURS OF INCUBATION 657 ventral aortic roots. Atthe cephalic end ofthe foregut the ventral aortic roots farn dorsad; curve around the gut, and then extend caudad as the pales dorsal aortae (Figs. A-20, A-24B and C, and A-25). The curved vessels ——ridge llantoie sak ore etized that the bursa of Fabricius is one of the major componen a i ‘mmune defense system of birds and that if it is extirpated, he : Chan’ Btd t0 produce humoral antibodies is sharply reduced (Gti wha Joped to conditions which urinary system is not yet developed 10 © arcane tose in the ‘adult, the parts oft which have been estblshee . alana sent associated with the cloaca. The, proximal pore” © opens di Stalk, which is the homologue of the urinary bladder fever retort ito the cloaca (Fig. 38). The ducts which drain the deve Stalk, Ty,0"82#8 also open into the cloacal region on either side ofthe free of thes stage lite indication of the formation of he BOM er embrygg wal UCAS to the cloaca can be discerned only by Felatio Dipindai dengan CamScanner580 DEVELOPMENT OF CHICK EMBRYOS FROM 18 HOURS TO~4 DAYS OF INCUBATION ui ont 7 Merencephaton as Ophthalmic dv, Ao.3teh i Maxie Mand, . ach F278 Hamby i enn oman arehT Fraune 4-40 The lower drawing is @ sight schematized represent Prarie! een ol w 3Y+day cick Ths secon i kay Dipindai dengan CamScannerDEVELOPMENT OF THE CHICK DURING THE THIRD AND FOURTH DAYS OF INCUBATION 681 ‘The Circulatory System The 4-day chick embryo possesses two extraembryonic circulatory arcs—the viteline and allantoic arcs—in addition to the intraembryonic. syste f vessels. The heart is already well developed. aie The pattern of the vitelline circulation in chicks of 4 days is shown in Fig. ‘Ax43. Blood from the dorsal aorta Teaves the body via the paired vitelline arteries, which branch repeatedly and end up as a large capillary network lying on the yolk sac. After picking up nutritive materials from the yolk, the blood flows into venous collecting channels and finally makes its way either into the marginal sinus or directly into one of the larger vitelline veins. The proximal portions of the main vitelline veins have fused to form an unpaired median vessel within the body of the embryo. Through this vessel, the blood returning from the vitelline circuit eventually reaches the heart. The allantoic circulatory arc is not yet highiy developed in 4-day embryos because of the small size of the allantois. The allantoic arteries arise by means of the prolongation and enlargement ‘of a pair of ventral segmental vessels arising from the aorta at the level of the allantoic stalk. Their size increases rapidly as the allantois increases in extent. From them the blood is distributed ina rich plexus of vessels which spread over the mesoderm of the allantois (Fig. ‘A-45). Later, when the allantois has expanded to meet the chorion as the chorigallantoic membrane, the allantoic vascular plexus serves as the main site of gas exchange for the embryo. ‘The blood from the allantois is collected and returned to the heart by way of the allantoic veins, which enter the body of the embryo through the allantoic stalk (Fig. A-42), course through the lateral body walls (Fig. A-41E to A-41G), and ultimately empty into the sinus venous (Fig. 44 aoe wi te intraembryonic circulation of the 4-day chick Begins with the ventral aorta, which leads into the series of aortic arches. Aortic ‘arches I and often II have disappeared as main channels, leaving only the third, fourth, and sixth pairs of arches. The right and left arches are s! symmetrical at this time. The regression of the first two aortic arches ploys & prominent role in the formation THaietuor aneries supplying the head--the external and internal carotid Seer are extensions of the ventral and dorsal aortic roots, respec- tae, originally pé J throughout much of their length, are by en fer core eal ate posterior pharynx (Fig. A-41B and C). As 4 days fused as far rds caudally, i gives off the paired vitelline arteries. Cen ae any traces of the unpaifed coeliac artery can be seen in 4-day Occasionally, caf Marther caudally, the pared allantoic arteries branch out chicks (Fit. A-44). Fig. Act). Throughout the length of the aorta, paired ee naan supply the somites and their derivatives, pmental arteries PR us ystem i represented principally by the anterior an inte inal veins, which drain the head and trunk and converge into the common cardinal veins as they turn medially to enter the sinus ven tag) The posterior cardinal veins ie just dorsal to the mesonephre hdres s Dipindai dengan CamScannerGanglion VIE & VINE anstion V earch of it. earotid ate a Anterior exiinal vein nt carotid artery Aortic teh I omic ach pharynx pene sree’ Mesocorle Dorsal ganstion Neural tube notochord ors ores Z Fan landibular arch clef TMandibul Srsinat Hyomandibula lets e ‘went Granehial groove I Mesococ'e ‘Sentory layer of retina Pigment layer of retina Tens Right common enira vn ‘ungbud Pleuslregon of cotlom 2 Lats commein eardal vein Vertile 82 . Pericardial resion of eoelam Dipindai dengan CamScannerDipindai dengan CamScannerDEVELOPMENT OF THe Cann CX OumIg 9 aren 683 INS TM mm an poyATH DAYS OF NCUBATIO Ductan helen Mesonephvie dey Dora! mesemery ~ Doral panetion ™ Assen ectodermal rite of Dome! mesentery, " srcerior soperdage bead SEL vets onephrc tole Altontov ein Dermateme wa F —tesoneghne ovet ben min { a ey en Posterior Dorta sore ~ tesonepheie Vitesira 6 uct Alantoic vein ves Hineget < Dons! son Noroenord ne tenes oo sraieaued on & canon of be secuons FIGURE A-11 ectons of 2 gay oH6% THE REE Diagrams of ransverse a ‘Small outline sketch of the enure eM no Situated to A-dIH). Situat | veins (Fig. ~ throughout their length (Fi i ove ‘ances! ventrally in the mesonephe ory portal system of more A-44), which are a relic of Dipindai dengan CamScannerFood materia mere i yolk se seiorbec in vlinevateulr £26 relove tery of the weste products riicent to their metaboliam. (From Patten, 1951, Am, Soventis, vo $6.) forms. The role of the veins in forming the inferior vena cava is discussed in Chap. 17 (Fig. 18-12). The early morphogenesis of the heart is summarized in Fig. A-46. By the third and fourth days, the heart has already twisted upon itself to form a loop ‘and the major gross divisions of the heart are recognizable. Expansion of the atria is prominent during this period. During the fourth day the truncus arteriosus becomes closely applied to the ventral surface of the atrium, which expands around the truncus and becomes subdivided into right and left chambers. Separation of the common ventricle into right and left ventricles has just begun ‘The Urinary System tis possible to recognize pronephric tubules in chick embryos of 38 to 40 hours (Fig, 17-24). In chicks of $0 10 $5 hours the mesonephrie ducts and primordial tubules are clearly identifiable just lateral to the somites (Fig. A-30D and E). BY the fourth day mesonephric tubules are well under way in their development. The metanephric tubules, which constitute the excretory units of the perma Rent kidneys, do not appear until considerably later in development, and the Benital organs which become intimately interrelated with the urinary organs are Dipindai dengan CamScanner685 eveLOPMENT OF THE CHICK DURING THE THIRD AND FOURTH pays OF INCUBATION FIGURE A-43 me cncuation in chick of about 4 days: he Lille) abbrevia- ne co teline voi, LV. Vara iSO To mate ‘nal vel (sinus terminals); PV. V, posterior vette ven, V. A, line artery, The drecton {blood flow is indicated by a1"o¥ 1 appearance. The later stages of the Fellas the entire sequence of stages in red in Chap. 17. also relatively late in making the vy system a development of the uninar the formation of the genital orgen' are cove The Coelom and Mesenteries cavity consists of three regions: In adult birds and mammals the body cof : pericardial, pleural, an es are paired, each of the id peritoneal. ‘The pleural cavi : : Dipindai dengan CamScanneral Vertebral artery Cervical segmental a Ante areh tv. Borie ach Vi Posterior eargina Cosine artery. Interior vera ex Subelavian vein Subclavien artery. Allantoie atery Mise artery FIGURE 446 Reconstruction of circulaory system of 4-day chick (rginal xs, reproduced x18). From the seme embryo #8 tnat represented in Fig. A-38. These itustratons should be susiedtopoties pleural chambers being a laterally situated sac containing one of the lungs. The the heart and the peritoneal chamber contain gs and heart are unpaired. These regions of the Raul body cavity are formed by the reshaping and partitioning of the primary body cavity, or coelom, of the embryo, In the chick the coclom arises by a splitting ofthe lateral mesoderm of either side of the body (Fig. A-47A and Bi), It is therefore at first a paired cavity, Unlike the coelom of some of the more primitive vertebrates, the coelom of te {he coelomic chambers are not limited tothe region i which the body ofthe Embryo is developing, They extend on either side into the mesoderm, which. in Common with the other germ layers, Spreads out over the yolk surface. Large Dipindai dengan CamScannerDEVELOPMENT OF THE CHICK DURING THE THIRD ANO FOURTH DAYS OF INCUBATION 687 FIGURE A-45 ‘The blood vessels of a 4-day chick. The basis of the illustration was the same wax-plate recon- ‘haction trom which Fig, A-44 was drawn. The smaller vessels and the primordial caplary plex ees taore added irom injected specimens. The labeled diagram of Fig. A-44 will sorve as a means of identitying the main vessels. (See colored insert.) tive coelomic chambers thus come to be extraembryonic in hap. 7; Figs. 7-1 and 7-4). The portion of the coelom that parts of the primi their associations (C id gives rise to the chap Te jody cavities is fist marked off by the series of i Ik (Fig. A-47C and D). fold: ‘h separate the body of the embryo from the yol! As frercisoreton Me ventral body wall progresses (Fig. A47E and F), the Dipindai dengan CamScanner6BB DEVELOPMENT OF CHICK EMBRYOS FROM 18 HOURS TO 4 DAYS OF INCUBATION Cephalic tp of "foregut ewe Font vin tortie root bend aid tet sore Endocardium —. ves Ameror a BX: A. 28 Hours 8. 20 Hours (8 somites) Optic vesicle = cepresion wenery Sroncus __ entice D. 3B Hour 40 Hous (ie somes) {1B somites) F. 42 Hours cht) Lens oteve Anseior Intestinal onal ©. 44 Hours (22 somites) (25 somites (29 somite) FIGURE 4x48 Me maj gratings of the heart and great vessels of chick embryos of vatiow Some of the major topographical features of the Wed asize the ro. sem ooeparhil eat ‘Abbreviations 20s have been outined to emphasize the relations umbere; Sin ven, sie vnosusy Wace en MUI IV, cric arches ol the designated Dipindai dengan CamScanner890 eve, “LOPME SMT OF cick Ewonvos Frou HOURS TO 4 DAYS OF mcusaTiON evra nate Noteehond Dorsal matodsem Somatopleure comte Endodeem Invermediate metoderm F ==> Somatie mesoderm Sa a Splanehnopleare Nevnitube ___ Sonite Spinnic mead Somatopiere ~__ Dor bora Imermedate mesoewm = Intrsebryericcolom ‘ae = Later! body fold € SS —— Sptancnoplese SS nreroryonie Primi pt = coworn Dons acna Somite Porstior sarin! vein A Meneaivi dct ad we (rom intarmadte messerm) Mecenchiyme Latwal aac tot Somstopleue Sotenemnapleure Primitive gut Neural robe Smite Masenehyme Doral sora Mesonepheot Borst mesentery —Z Bottom Gast. Liver in: hepanie J Pspeeating ventral ‘omentum ventral mesentery Ventral mesentery Figament 6 of liver ‘i Right and lek AQURE aay coelom content jematic diagrams of cross sections Dipindai dengan CamScannerJed bel ers of splanchnic Me de onstitut pe mesial walls of The iat and lett coelomic rs, The double ayer o splanchnic mesodt which thus becomes 21 sed to the gut and ypport it site body cavity Hh Tynown as the primary ™e enter. THE if th mesentery dorsél tO the gut, suspending 110 ijorsal body Wall, 8 e Mersal mesentery: 3nd ert ventral to the EH sai to the ventral ‘body wall, is the ‘ventral mesentery. ey en the dorsal and eeearal mesenteries Te jars, they conse complete membranans © ition dividing THE vy cavity in and left ree. The primary 4 eet mesentery persists cage part, but whe YEATES mesentery soon extensively resoroe 5g. AXATED) inging ght and left coelomi chan int onfiuence ¥‘ ntral tO whe and ¢st splishing & 1e ‘onpaired condition ody cavity Mraraceristie OF tHE A are ration t0 1 ey nesenteries OF IRE the Tesocardia may DE regarded 25 SP nT egions of the VERY mesentery HE ‘post cephalic Pa of the b ‘cavity, th ties embed ied in the 60! ‘pody wall inste of Peing suspended OY al mesentery 2510S sei ety (F.Fies: A nd A By gk mesentery Y Sever, GEVEIOPEA annet “anteriorly post only, and whe whe heat" yemed it 5 suspended if he most 2 part of thi twat meseneny: Ty sor 2 vent : arts of tHE esocardia Me gorsal t ee Je heart ectively (Fis esenneny tral ,gocar dium tle tater th sal mesoc’ pene a escent ant an el ‘emi x come fluent BN aeicardal i ay cast 25) Di ipindai dengan CamScanner f