QL *c JH 15 s EXCHANGE BIOLOGY LIBRARY G LIBRARY Inn- re/inn ft 'I j'mm Cailiard's Medical Journal, _ hy >,,, m • in tli, /,rnj, MBTH <'tBccum»s IN THK EMBBYO: PLAO FFLE THE Ax.UX" IlESPIKAT BKPIRATIOX ix THK XEW-BOBX : THE CHANGE is MECHAXICS wmcn THB YI58 TO THE AlB-CHAUBEB IX THE EGO. \I Di THE TISSUES Or THE WoMB.* Br W. H. TKII'I.riT. .V I M seen that respiration and circulation most be treated together, since i a connected movement for pumping the commerce in and through the organism for elaborating structure and evolving force, and ***** the whole • founded in the power of producing rhythmical changes in pressure, the fluids flowing from high to low pressure in conformity with organic law ; that the ••stines and bloodTeasds are necessarily allied in respiration in order to make importation and transportation a connected movement between the ceD-timod and environment, which the scheme calk for. In other words, that xstptM? action in respiration compel* the commerce in the Teaauh, while the action in the heart, arteries and venous system circulates it through the organ the action being unified throughout by means of the nerroas com- binatians in the medulla oblongata, in which the respiratory, raso-motor dontary motor centres, are correlated, the one calling for the other, as has folly set forth in the preceding pages, and that the vital phenomena, :nical and physiological, appertaining to respiration, circulation absorp- tion, etc., are readily explained and accounted for, giving absolute proof of the correctness of the premises, as before remarked. We now follow this matter a little farther, and take op CI BrrLATIOX IX THE EXBBTO, a subject which, it most be admitted, is at present veiled in deep obscurity. This is nevertheless susceptible of ftrjJhmmtitw^ The embryo is an aquatic •""" * — — y "•"*• *-'~ri — I ' *~ r-"— "— — •"**-* "~- — T ruiinnaij if tii Body : or. tfte Xeehaoic* in Scepintjoo. airrtrttM Absorption. Etc.,- ehapun zrt Md rrii. r :. : /.:"•:••/: : •• • « ; •; ... animal, since it leads a subaquatic existence, being placed under water (Fig. 1) and deeply buried in the maternal tissues, which would account for the peculiarities that obtain in its circulation approximating it to the stages in de- velopment represented in fishes and amphibia. FIG. 1. — GRAVID HUMAN UTERUS AND CONTENTS, showing the relations of the cord, placenta, membranes, etc., about tbe end of the seventh month. 1, decidua vera ; 2, decidua reflexa ; 3, chorion; 4, amnion. (After Dalton.) The physical conditions under which the embryo is evolved determine the special vascular arrangements for effecting circulation, while the trans- formations which accompany this provide for the radical physical changes which are ushered in at the end of the term, when it becomes an air- breather, the fundamental circumstance underlying it all being an adjustment ivith pressure, and the power of effecting rapid rhythmical changes in pressure, or of producing the universal pumping actions going on in the body. For ex- ample, we have seen that the rhythmical expansions and contractions pervad- ing the body in the air-breather, and known as respiration, compel oxygen and aliment in the circulatory apparatus for evolving force and producing growth ; so, in like manner, a similar necessity exists in the embryo for compelling the nutri- tive and force-producing elements in its circulation for producing growth and evolving force, which is principally expended in elaborating its structures. But since the embryo feeds in the uterine sinuses from which the com- merce is obtained, and into which the waste products are returned, this calls for the differentiation of the special organ known as the placenta, and which answers to the more highly differentiated lungs and intestinal canal which are to substitute it at the end of the intra-uterine term as the relative adjustments with the larger environment, lower pressure, and higher order and amount of work which this involves.* The placental souffle, then, which is distinctly heard through the maternal structures is the analogue of respiration in the air- breather, the relative ratio of the movements to the pulsations in the foetal heart •For further particulars, see Part II. of this work, "On the Relation of Gravitation to Development," to be issued shortly. being also the same, or as 1 to I of tin- latter, whilst the villi are the analogues of tilt* villi is tin- intestinal canal, tho ono 1n'ing submerged in the sinuses, the other in the juices in the intestine. Since the pumping action in respiration is absolutely es-ential for compelling the comineive in tin' vessels, it follows that this circumstance shmilil l)o represented in the t>mbryo,\i8 the maternal blood (loos not enter the embryo, the latter feeding out of lEe sinuses simply by menus of the villi in the pkoenta/and tliis pumping action spoken of ; for here, as elsewhere in the Ixxly. there are no means for increasing circulation but by rlii/tliiiu'rtil I'/HHt'ifx in fi-is.-iii-t'. The result must then be the action in the plaeeiita simulating respiration. The relative frequency of this movement to the action taking place in the ft-d miiM-l. ^ are present for hflMMillg tin- .•iiiT.i.'v nf tin- rhythmical expansions ami c. mil-actions. But muscles arc nut essential in this action, a notable example nf which is bnythed in tin- In-art itsi-lf, which expands ami cniitracts rt'gularlv and rhvth- micallv liefon- a nniscl.- or m-r\.- is ili-\ -loped in tin- protoplasmic and un- diH'en-ntiated \vallsip. -J'.l'.li, also in capillaries ami in earlv animal forms. It is interesting to note tin- fact in ttris connection i which has already been comim-nt^l upon in tin- higher stages of ili-vdnpnicnt i. that <-r,ri/ ///<»(•.•;/<«•«/ of the animal, l.v rxtrinHni,' nr retract in.i; tin- mantle ami tin- Imincln-il ]K3dtlLcle8, must ./'// increase circiilatimi ami n-spiration in roiTespnnilence. It could not In- ntherwi>e. Niiim-r..n- -tminita in tin- rounded extremities of the peduncles (a, a, "i atl'onl fi-ci- in^i-i-ss mid ogress to the tlimls in the central canals (b,c). Fin. 2.— MF.DI-SA (Rhtinntoma cuvieri). A, a, a, eight peduncles; 6, c, internal canals leading up to the stomach; trillions separating the caviiy of stomach from the ovarial sacs; i, i, external opening to ovarial sacs; m, oesophageal passage. (After Grant) The capillary loops at the sides of the peduncles (/, k), which function as ab- sorb-in; and respiratory vessels, may be taken as the simile of the capillary hxips in the tufts of the placenta. This brief description will serve to explain the mechanical principle in the placenta, and it now remains to be seen how this action is assisted by the action takin j place in the womb itself ; or how the maternal and foatal circulations are made to connect for effecting mutual interchange, the former supplying the nutritive and force-producing elements for the growth and elaboration of tissue, the latter yielding up the waste products to be borne back through the maternal channels to the environment from which everything is derived, and into w!iich, in due time, everything is returned in the form of waste and tin il dissolution. MODE OF CONNECTING THE MATEENAL WITH THE FCETAL CIRCULATION. The circulation of maternal blood in the uterine and placental sinuses is the same in principle as that for circulating air in the lungs, namely, by rhyth- mical changes in pressure in the sinuses, which is produced by the action taking place in the womb itself ; taking the placental sinuses to represent the alveoli, and the uterine the tracheal system, the blood flowing in and out of this system of canals by^reflux action for renewal, just as the pulinonic air flows in and out of the tracheal system for renewal, only that in the former the fluid passes into the venous system of the mother and is returned by the arterial, both terminat- ing by capillary openings in the canals, while the placental souffle answers to the respiratory murmur. We have already spoken of the role which the placenta plays in this respect, and it now remains to take up the very important one which the uterus itself FIG. 3. — LONGITUDINAL SECTION OF A LATELY GRAVID WOMB, showing the canal system. Taken from a case of post partem bffimorihage (partly diagrammatic). 1, peritonieum ; 2, uterine canals (sinuses); 3, openings to canals on the mucous surface corresponding with openings to placental sinuses; g, Fallopian tube; /, ligament toTtrtCTyr-r, round ligament. (fir-tv^^l performs for completing this deeply-interesting mechanics, which it would be difficult to overestimate, underlying, as it does, embryonic life. First, with regard to the uterine sinuses, we may mention that this canal a which is channeled in the walls of the gravid \\,m\l) does not consist of veins or arteries enlarged for the purpose, but, on the contrary, that the proper substances of the womb itself (muscles and connect he tissue i compose the actual walls, which are lined by a tleliente uieinbnine answering to the lining mem- brane of the vessels. The difference is very material, since the vessels are enabled to expand ami contract themselves for increasing circulation, while in the ca>'' of the sinuses, the walls of the womb would have to expand and contract in order to HIKW«- circulation in them, mid that this is the case will appear more obvious as we proceed, the mechanics undoubtedly calling for such a condition. When a longitudinal section is made of a gravid womb at full term, this canal ^v-teni is seen to traverse its substance in various directions from near the central IM nt ions, but trending to the internal or mucous surface where they connect with the placenta (Fig. 3 — '2, 3). They never penetrate to the external surface to form continuous tubes with the arteries or veins; but on the contrary, both these systems of vessels connect with this system of canals by means of capil- Itirit ••*, the arterial emptying and the venous discharging from these common mains. Hence this blood is mixed as a matter of necessity. The next important point is the method this system of canals exhibits in con- necting with the placenta! sinuses, which is done by projecting the lining mem- brane in/') l/i'' t>>(lixtan<-e of the placenta (Fig. 4, c, c, c, c). This spreads out at once FIG. 4. — VKIUII u. SI:CTHI\ OK I'I.ACF.NTA, showing arrangement of maternal ami f, >>, ileciilua; 0, < . . . . , urillres of uterine sinuse.j. (After Dalton.) to form the large cavitary spaces in which the highly ramified placental tufta are suspended and submerged in blood. The walls of these large cavities in which the lining membrane of the uterine sinu>es is merged answers to the outer surface of these highly-rami- fied tufts which tit into this as the fingers in a glove, the vascular loops being contained inside of them (Fig. 5, a, a). It is easy to perceive that this out- side wall and the wall of the capillary itself in// rr< n<- lx>tween the. maternal and foetal blood, and that interchange would have to be effected through these two. 8 membranes ; hence, to be very active it would call for considerable force for producing this. FIG. 5.— EXTREMITY OF FCETAL TUFT, from human placenta at term, in its recent condition. A, a, capillary bloodvessels. Magnified 135 diameters. (After Dalton.) During attachment, when the sinuses are filled with the maternal blood they are greatly larger than after detachment, when the blood is forced out, and the tufts are compressed against each other in the maternal passages, reducing the placenta fully one-half its size. At the time of birth, the necks of the uterine sinuses, where they join the placenta, are readily torn through (Fig. 4, o, c, c, c), and, peeling off easily with the decidua, the organ is expelled, unless, forsooth, abnormal nutritive changes in the parts have made the union more intimate. The problem in the mechanics concerns the manner by which the blood is speeded in and out thesf sinuses commensurate with the increasing wants of the growing embryo, and how this in turn is connected with the arterial and venous systems of the mother for compelling correspondence in them, the whole forming & connected movement — a movement within a movement, so to speak. We have seen that change of pressure is the law of the animal circulation., ;md it now remains to make rigid application of this principle in mechanics to the special phenomena, .anatomical and physiological, appertaining to this inter- esting region, in order to make them also intelligible, which otherwise are utterly inexplicable. It follows, that for increasing circulation in the placenta we must provide for rhythmical changes in pressure, since no other force applies for producing it, and we must connect the action in the womb (which expands and contracts regularly and rhythmically) with the action in the placenta, so that when the latter [expands for producing low pressure, the former contracts for producing the opposite, and vice versa. In this manner an active circulation in and out the placental sinuses could be readily produced. The following facts may be given in support of this opinion : 1. The womb must expand and contract regularly and rythmically in order to increase circulation in tin' sinuses. since tliis is essential for changing piw-.ure upon tin- l)l walls of the womb are the provision for more energetic action than is possible to the placenta, which is composed almost entirely of vascular loops. '2. The womli .-.ii IT* HI in/* -the emliryo. and its a<-tion would therefore be more etl'ective for producing the changes in pressure upon it, which is also in imilnlii •/. <•(' ir/nit t'iki'x /ilni-i in /In- uiitiiinn of the grow ing chick, it beJBgZOOked to mid fro in the egg b\ the slow rhythmical expansions and contractions taking place iu the amnion i|i. -I'.I'.M. and by alternating this ai-tion with the one taking place in the placenta, it is readilv jH'rceived how a rapid circulation could be main tained in the maternal and placental sinuses for compelling correspondence lietwecn this and the energetic circulation in the emliryo, since it all forms a connected whole. This would also be in conformity with the principle in the circulation, the blood flowing from high to low pressure. As the placenta i-.i-fiiimlx for reducing pressure within itself, and for increasing pressure in the emlir\o. *///, elliiig this blood in the placenta, at the same time by increasing pressure upon the embryo it should determine a more rapid movement from the latter to the placenta ; the one involving the other. But when the placenta contracts for increasing pressure (thereby compelling the blood out of itself in two directions, or toward the embryo and the maternal sinuses), the uterine walls expand for reducing pressure in the embryo and the uterine Minist s, thus greatly expediting the placental efflux; and, taking it all in all, there can lie very little doubt but that the force which is represented in the muHled murmur of the placental souffle is mainly the product of the muscular uterine walls, though l>oth undoubtedly contribute to it. '.'<. This action in the uterus and placenta would account for the very curious and suggestive n/i/ii/n//// in that portion of the canals connecting the uterine with the placental sinuses, being upon a line almost paraUeJ with the transverse axis of the uterus and placenta, which is precisely what is called for by the special mechanics, in order to effect the lateral or to-ond-fro movements, the vessels simply elongating and contracting with these, as the case may be, with- out interfering with the calibre of the tubes. Were the vessels straight, it would be utterly impossible to operate this mechanics, since the lateral move- ments should obliterate the vessels by closing the calibre. It could not be otherwise, in the very nature of things. In the accompanying diagrams (Figs. 3, 4), the vessels are represented as nearly perpendicular, but this is done simply for better definition. I. lint the strongest proof, perhaps, of this higher function of the womb is furnished in the \asmlar and nervous connections subsisting lietween it and the maternal organism. And as all this relates to circulation for building up and elaborating the embryo, obviously the mechanics for increasing circulation in the womb commensurate with that in the embryo should extend to the vessels of supply or the feeders, as also the discharging vessels, or the arteries and \eins. It must be shown how this new movement, this new life set going 10 within the other, twines its arms around the maternal vessels and feeds itself in the measure of its necessities, by means of this pumping action in the womb and placenta which represents respiration. The spermatic and uterine arteries are the feeders (Fig. 6, OK TI1K NINTH MONTH. .1. tin' fnndus and bod) o( Ilu- uterus, having Hie peritonaeum diss, left bypoaMlta nerve B left hjpou'astric ^iingliii, and ^ivin^ oil branch'- to the left gub-peritoneul ^an^lia : ,i. hxmorrfaolda] nerves and arterj : •/. sacral nerves entering the whole outer surface of the hjpoga-trir gunglion: r, left hypogiistric ganglion with the arteries injected; »•. nervi- of i '. nerves with an injected arlen proceinling from the upper part of left hypn- L':i-tric ganglion along the body of the uterus, and terminating in the left spermatic ganglion: 'i. continuation of these nerves and the brunches which they L'ive off In the snb-peritoneal plexuses: /-. >oine nen es passing upward beneath the sub-peritoneal plexuses; and an nici.-ing fiee|\ with them; "•. left :-pcriu:itic irangliou. in which the nervea and artery from the hyiiog.istric ganglion, and the branches of the lefi sub peritoneal plexuses terminate, and from which the nerves of the fiindus uteri are supplied: .r, left sub-|ieriton«'al plexuses covering the Ixxh of the uteru>: //. left sub-peritoneal ganglion, with numerous brandies of nerves exten. I ing between it and left li\ nerve and ^arglia: ^, left common iliac artery cut across and turned aside to expose left hypogastric nerve and ganglion. (After Dr. R. Lee.) 12 and if this means anything it means that circulation shall be in correspondence with the physiological requirements, or supply equal to demand, the cardinal circumstance being the f the embryo, which is the object and purpose of the organ, the others being simply incidental. It conies to this, namely, that the nervous force for expanding and contracting the uterine sinuses should expand and contract the uterine bloodvessels at one and f/ie ner of muscles and nerves in the woml) are for compelling Otti the cmiteiits at the end of the term simply. Imt, on the contrary, that they jH-rform an active and essential part in the work of construction whirh ]>rci-edc8 expul- sion, lieyond a shadow of a doiilit. they arc not idle in all this while. :ally when supreme necessity would h:ive it otherwise; Oil the contrary, they an- n* /'<•/•'- /•«/ for carrying on circulation, while at the end of the term they are available for insisting in expelling the embryo, hence pe, form an active ;••./< from the beginning to 'the end of their existence, as is e\er the c;i.se with tin- miiM-les and nerves. ( )nc oth'-r circumstance in tliis connection, namely, the very l i, which undoubtedly is adjustment with this action, |K'rinittiiif; the rhythmical ex]iansions and contractions to take place without involving any strain to the vessels. The veins, it will be perceived, taki > a straight (Miu-sc, wliile the arteries are serpentine or ln>nt njvm themselves. HUB is tine to the fact that the veins are more extensile, and possess greater IMI-.VCI-S of elon^atiiiL,' and shortening than the arteries, the yellow elastic coat of the latter tending to limit their actions. It will IK? remembered that this cir- ciini.-t nice has forcible illustration in th" splenic artery and vein, the former being almost twice- the length of the latter, to allow for expansion in the stomach when fixxl is taken, otherwise this would involve prodigious strain to the vessels, with great re luction of the calibre. But the same remark will apply to the ves- sels of all the hollow viscera. Thus everything is in correspondence. Of course, the arc of movement in the womb and placenta is necessarily more limited than in the lungs, in 'which considerable space is required for sucking in the air simultaneously with the venous blood, but which would not apply for the foetal circulation, as the oxygen is furnished by art.-ri.'il blood at "/»• «ml the samr lini' with the nutritive and force-producing elenvnts, which the scheme calls for in order to generate force, since it is by a combination of the two that force is evolved. The inclusion of the intestines with respiration by means of the I meumogostric nerves has its explanation in this circumstance, as has been fully set forth in the preceding pages. Differentiation in the organs cannot, for obvious reasons, work any change in the fundamental principle underlying the mechanics for increasing circulation, which is by rhythmical changes in pressure involving a pumping action for compelling the commerce in the bloodvessels and expelling waste products, while the speed of the currents thus produced is determined by the rapidity and energy of the rhythmical ex- 14 pausions and contractions pervading the organs, inclusive of the heart ami vessels, since it all forms a connected movement for increasing circulation be- tween the cell brood and environment, from which everything is obtained and into which the waste products are returned, the two going on simultaneously. With the expiration of the intra-uterine term, expansion of the maternal passages sets in for reducing resistance to the egress of the embryo, and the womb and abdomen contracting simultaneously for increasing pressure in the womb, the contents are compelled out in the environment. Here, as elsewhere, the Jaw of pressure applies for compelling n/nri'/ncnf in flic con- tents of the, Iwlloiv viscera for which xj>friut tlir contact nf tin- sentient surface \vitl> the stiiiuilus in tho atmosphere itself woiiM also excite it. And if tin- child should lie injured by tin' rude experii-ne.-s incident il to |>.-irtiiritii>n. a yet more powerful means fur exciting tin- reflex actions connected with respiration is furnished by the su.ldeii application of cold t i the surface, as in sprinkling cold water upon it, or a sudden, shurp slap with the open hand may lie substituted instead, as is commonly practised. List, lnit not least, c.irlionie arid, ;LS it accumulates in .the blood, acts UK a special stimulus to respiration. It cries out hi pain, and, presto! the liemal mechanics is changed. The low pressure which is produced in the aheoli by expansion of the lung's during inspiration compels xiiiinllnin-onn afflux of air and blood in the alveoli ; while the high pressure which is produced l>v tin- subsequent c-int ruction during e\]iinition causes *////«// efflux in these tluids. which (low from high to low pressure in conformity with organic law the one (lowing out liy retlux action through the route of ingress, the oilier p.-is.,iiu into 'he left chambers of the heart and arterial system on its way to the cell-liroo 1. as has already been described in the air-breather. This •I'.aiidonment of the old route for the new is readily explained, since it is in sti ict accordance with physical law, l>eing in the direction of least resistance. For example, we /<•:>! r. ,ui'l the glcin an Tun- ia of • •.!-> 'li'inninlratiiMi in tin1 tulult liy tin1 sann- IIH- i;is. Kur fxample, every impact of mid walrr • tin* akin i>r."'l !.••'•- <|i i~ n<> lir iuspinilinu or ex[> union ill tlie luiigi ; not deep, however, l>ul very energeiir. 1(5 poured into the right auricle, that from the upper cava passing at once in the right ventricle, while that in the lower cava (which includes the blood from the umbilical vein) passes through the right into the left auricle, with which it directly communicates, guided by the Eustachian valve, but also pushed over and deflected in this direction by the weight of the descending current from the upper cava ; but if the head be downward then by its own weight the blood would gravitate in this direction, the influx of blood from the upper cava also compelling it. And 'with the absence of blood as a counter-force in the left auricle this blood is necessarily compelled in the latter, thence in the left ven- tricle and aorta, while that in the right ventricle passes in the arterial system at the aortic arch by way of the pulmonary artery and ductus arteriosus. After birth, however, the pumping action in the lungs reverses all this in manner as above described. The pulmonary artery in the embryo, in place of dis- charging through the lungs, left auricle and ventricle, empties its blood at once into the aorta as it passes under the arch, and which is also in the direction of least resistance, since it is impossible for this blood to thread its way through the capillary meshes of the unexpanded alveoli. This circumstance has forcible illustration, even in the air-breather, and when the alveoli are filled with residual air, by simply inhibiting inspiration by closing the mouth and nose so as to prevent expansion in the lungs, the blood, in consequence, rapidly accumulating in the right side of the heart and venous system. Tn the space of a minute there is lividity of the lips and whole cutaneous surface from venous stasis in the systemic capillaries. If longer than this, an appalling venous suffusion pervades the surface ; in the face most, for this is the most vas- cular portion, with the large venous trunks in close proximity to the heart. Even the eyes are forced outward, becoming prominent from distension of the iutra-orbital veins caused by obstruction in the cavernous and lateral sinuses. But the instant the obstruction is removed and the lungs are permitted to expand the dammed-up blood surges through the alveoli, and all runs on as be- fore. In other words, the heart and vessels are unable to carry on circulation in the absence of the pumping action in the lungs, for which afflux and efflux of air is essential. But all this has been sufficiently explained. INCUBATION. CIRCULATION IN THE EGG. Why should there be an air-chamber to the egg (Figs. 8 and 9) ? We are now prepared to furnish a scientific explanation to this physiological problem, otherwise inexplicable, viz., the contents of the egg for developing the chick is enclosed by a firm unyielding wall of living marble, and since the animal circu- lation is dependent upon rapid rhythmical changes in pressure, it follows that provision should be made within the shell for effecting this, otherwise the action in the heart and vessels could not take place. This air-chamber (g), together with the important relations it sustains to circulation and elaboration in the growing chick, organilogically, therefore, must be regarded as one of the most essential and important elements in egg-structure, the underlying principle to all the nutritive changes which are ushered in under the action of external temper- 17 iituri'. The Hccomiunvin^ illustration i FiU'. Hi will serve to impress the mutter. Tin- discipline in tin- nutrithe processes requin-s the blood to be brought from tlie vitellus and a • rated in the allantois. thence to be dispatched through the Imdy territories. Accordingly tun great venous trunks (omplialo-meseraic veins i, one in each fold of the aplanahnopfetm, embnofng the vitellus, are the KI<;. s. MuiiRAM OK l-ntti.'- Koa. A, yelk; '. vitel!im> membrane; c, chnlaziferous membrane; d, albumen; e. f. miililli* anil internal shell memiirane.-i : y, uir chamber; A, calcareous shell. (Aft«r Dalton.) first evolved, while at the terminal ends or confluence the heart is formed by the blending of the walls of these venous trunks. According to His. the heart is developed by the coalescence of a layer of the splauchnopleure with a similar layer from the somatopleure, the hollow cavity formed by the union bein^ in free communication with the adjacent omphalo- FIC. !). — Eoo or FOWL is I 'HOC BSD circulation, terminal sinus, etc. [)KVKt.opjiEXT, showing area vasculo.ia. with vitelllne meseraic veins. According to Foster and Balfour " the upper end of the heart is developed out of the mesoblast of the splanchnopieure," but " increases in length step by step at the expense of the continually coalescing ouiphalo- 18 meseraic veins." Thus the fact is undeniable, that the heart is developed in the venous system in connection with the vitettus. The rhythmical ex- pansions and contractions in this organ (the punctum saliens of early authors) serve to pump the vitelline fluids in the bulbus arteriosus and the two aortse which are developing at the other end of the heart. But for this air-ciishion within the egg (Figs. 8 and 9, g), neither these rhythmical expansions in the heart nor the changes in pressure for compelling circulation in the vitelline fluids could take place, since the unyielding shell would inhibit these actions, as must appear obvious. Respiration is provided for in the following manner : The aUantois (a diverticu- lum of the intestinal canal) is pushed out around the amnion which contains the embryo, and expanding its vast capillary net-work of vessels (whose footstalks spring from the two iliac arteries, as do the umbilical arteries in the mammalian embryo) against the shell-membrane or chorion becomes the respiratory organ of the chick, by means of which the venous blood is constantly arterialized, the oxygen passing in and the carbonic acid passing out through the pores in the shell by the action of the polar forces. But " at the time the heart first begins to beat, the capillary system of the vascular and pellucid areas is not yet completed, and the fluid which is at first driven by the heart contains, according to most observers, very few corpuscles The course of the blood then, during the latter half of the second day, may be described as follows : The blood brought by the omphalo-meseraic veins falls into the twisted cavity of the heart, and is driven thence through the bulbus arteriosus and aortic arches into the aortic trunk. From the aorta by far the greater part of the blood flows into the omphalo-meseraic arteries, only a small amount passing on into the caudal terminations. From the capillary net-work of the vascular and pellucid area into which the omphalo-meseraic arteries discharge their contents, part of the blood is gathered tip at once into the lateral or direct trunks of the omphalo-meseraic veins. Part, however, goes into the middle region of each lateral half of the sinus terminalis, and there divides on each side into two streams. One stream, and that the larger one, flows in a forward direc- tion until it reaches the point opposite the head, thence it returns by the veins spoken of above, straight to the omphalo-meseraic trunks. The other stream flows backward, and becomes lost at the point opposite the tail."* The following from the same authors is deeply suggestive: "Soon after its formation the heart begins to beat, at first slow and rare pulsations, beginning at the venous and passing on to the arterial end. It is of some interest to note that its functional activity commences long before the cells of which it is com- posed shoiv any distinct differentiation into muscular or nervous elements." It would be difficult to overestimate this circumstance, since it establishes beyond peradventure the power in the higher as in lower animals to effect rhythmical expansions and contractions in the soft tissues in the absence of any muscle or nerve for producing them. But at present the significance of these rhythmical expansions and contractions taking place in the heart concerns us most, since the manifest purpose is to increase circulation ; and as this can only be done by * Fos'.er and Balfour's •' Embryology." 1'.' pumping the blood of the omphalo'iMSwaio veins, it follows that both expan- sion and contraction is necessary for accomplishing this, the one for aspirating this fluid, tin- other for pn>|>ellini; it. This would explain why the action should In-ill a placed ii]x)ii it ; notablv, the former is produced \>\ tin1 ainniotic fluid and the latter by the action in the ainnion. Tin' aiuiiion closes around the embryo of the chick on the fourth day, and on the fifth tluid U^'ins to collect in the sac, and by the seventh the embryo is submerged in a considerable quantity of water. "By the seventh day very obvious movements U'gin to api>ear in the amnion itself : slow vermic- ular contractions creep rhythmically over it. The amnion, in fact, begins to pulsate slowly and rhythmically, and by its pulsations the embryo is rocked to and fro in the egg. This pulsation is due, probably, to the contraction of involuntary muscular fibres, which seem to be present in the attenuated jHirtion of the mesoblast, forming part of the ainniotic fold" (Foster and K-llfoUM. The physiological significance of this accumulation of amniotic fluid, and the rhythmical contractions and expansions in the amnion, may not be doubted for a single moment, since the former would increase pressure, while the latter should produce the necessary changes in pressure in the embryo for compelling respiration and circulation to be in correspondence with the nutritive and func- tionid processes in the growing chick, both of which are constantly extending their limits and requiring more and more force for effecting them. These slow pulsations in the amuion of the chick answer to the placental and uterine souffle in gestation, the principle being precisely the same. How other- wise explain this circumstance ? But, as has already been remarked, a II pulsa- tion* relate to changes in pressure, and these pulsations in the amnion, together with the amniotic fluid, relate to changes in pressure in the embryo for increas- ing circulation of the juices. The explanation of the mechanics is sufficiently easy ; notably, there are two cardinal points from which to regard it — one in the allantois, the other in the embryo. First, commencing with the movement of expansion in the allantois. The increase in pressure which this produces in the embryo through the am- niotic fluid should cause the venous blood to flow with increased energy towards the allantois, the point of low pressure within the egg (the heart and vascular >\ stem, of course, assisting in this); the contraction or condensation of the amnion. by relieving pressure in the allantois, enables this to expand part passu with contraction in the amnion for aspirating the venous blood, at the same time that it aspirates the air through the outer membrane and pores of the shell. When the movement is reversed by expansion of the amnion, the reduction in pressure which this effects in the embryo, together with the simultaneous in- crease of pressure it produces in the allantois by forcibly compressing this against the shell wall, causes the aerated blood in the latter to flow with aug- mented speed into the heart of the embryo, as also through the tissues of the 20 1 'liter; since thet blood vascular system woiild be less embarrassed and be more free to act in cousequei:ce. . |But the allantois itself also participates in this action, the fluid it contains enabling it to effect such rhythmical compression of the capillary plexuses TIG. 10. — DIAGRAM OF YOUNG EMBRYO AND ITS VESSELS, showing circulation of umbilical vesicles, and also that of allantois, beginning to be formed. (After Ualtou.) (Fig. 10). The following forcible illustration (Fig. 11) by the distinguished biologistjat Jena will serve to impress the matter. At this early stage in develop- ment (third week in gestation), it will be seen that pressure is increased at the •cardinal points, namely, vitellus (a), the body of the embryo (c), and allantois FIG. 11.— HUMAN EMBRYO IN THE THIRD WEEK. A, large globular yelk-sack; f>, allantois; c, amnion ; d, tufted cborion. There are yet limbs. (After llueckel.) (b). As the embryo and allantois are elaborated out of the material in the vitellus, this would explain the greater accumulation of fluid in this locality for compelling circulation toward those two points, whilst the rhythmical contrac- tions of the yelk sac should greatly expedite it. For increasing circulation •Jl between tin- cnibrvo :uul aliantois (c, /'i, commensurate pressure is prcxluced by iici-iiinuliitiiHi of fluid in these two jitiints or jioles of tin- circulation. This, to- gether witli tin1 action in tin- mtmihranes themselves, and the heart and vessels, is siillicient forcaiTvine; on circulation in the iaitiul stages of embryonic evolu- tiou ; lnit witli tin- incr. "\\tli comes increasing difficulty for effecting it; hence the pumping :ictii>n whicli is set up in the placenta and womb, together with tin' accumulation of iimiiiotic lluid for transmitting these actions upon the embryo, a.- described above. Tlnis, even tiling is in correspondence — the liquor iimuii, the inoreaoBg growth of tin- placenta .-mil the number of muscles and nerves in the walls of the womb — and so continues till the close of preg- nancy. In other \vonls, it all forms a coniu-ctc d whole in the mechanics of the i-mhryoiiie circulation. In tlic case of the bird, the jillantois answei-s to the placenta, since it pumps in both o\\L,'en and nutriment ; only that the pumping action in the abdomen (the soft hinder parts of the bird) for pumping air and blood through the alveoli, is set up in the hitter days of incubation, when rapid atrophic changes soon obliterate the umbilical vessels, and, breaking the now attenuated and fragile sht 11-wall witli its b«ak, it finally makes its escape, leaving the allantois and atrophied membranes behind. MODE OF GRAFUMi Till. o\l\l l\ THE TISSUES OF THE WOMB. In jiHiiiiiH'iliii tho ovum is not discharged from the maternal passages, but i« retained in the expanded oviducts at the point ol juncture, which answers to- FIG. 12.— A VERTICU Sn Tins FROM THE FIG. 13.— Same tubules, greatly magnified. UTERINE Mi'cors MEMBKAXK, showing the (After Dalton.) numbers and position of the tubules. A, free surface ; 6, attached surface. (After Dalton.) the womb, where growth and elaboration are effected through the vascular con- nections established between it and the maternal circulation. Briefly, the mode of closing this is as follows : The uterine mucous membrane is virtually but a dense mass of single, straight follicles, arranged perpendicularly to the free sur- face (Figs. 12, 13), and with the closed end resting against the muscular 22 walls of the womb. At intervals, fine bloodvessels course up between them, and, reaching the mucous surface, surround the tubules with a capillary network. But when fecundation is effected vascular turgessence at once sets in, producing rapid growth and development of the membrane — of the follicles especially ; and, coming in contact with this highly vascular and tumified surface as it leaves the Fallopian tube, the ovum is rapidly incorporated with it, by means of the •shaggy villosities which are thrown out from the outer surface (Fig. 14, «..), FIG. 14.— OVUM OF THE RABBIT, from a Graaflan follicle J6 of an inch in diameter. A, epithe- lium of the ovum; 6, zona pellucicla, with radiating striations (vitelline membrane); c, germinal vesicle; rf, germinal spot; e, vitellus. (After Waldeyer.) FIG. 15.— IMPREGNATED UTERI'S, showing connection between villosities of chorion and decidual membranes. (After Dalton.) which grow into the expanded orifices of the tubules, and function as temporary villi for pumping the abundant albuminous secretions of the glands in the in- terior of the ovum. Of course, they contain all the elements of tissue struc- ture. With the ingrowing of the shaggy tufts of the chorion in the follicles, the mucous membrane expands around the ovum for bringing the follicles in contact with every portion of the chorion till the whole is completely enclosed, forming what is known as decidua reflexa, while that portion of the membrane •with which it first came in contact, in immediate relation with the muscular walls, constitutes decidiia, vera, the part that enters the structure of the placenta which is subsequently formed. At this time the villosities of the chorion project into the uterine follicles in every direction (Fig. 15), but as growth proceeds, aiirl more and more nutriment is needed in the embryo, the villi in relation with decidua vera become vascular, while those in relation with decidua reflexa are atrophied, in consequence of which this portion of the •chorion becomes bald. Finally, the transformations which occur in the placenta ' * • convert the tubules int.. pl.-iccnUl sinuses, and tli.' villi of the chorion into vaa- culiir loops, •Demanded and embraced liy the lining membrane of the HiiiUKOM, like the finders of a glove i I'i^. 16), aa before remarked; hut since the osipil- l.irv lm>ps ore projected ill the villi, the natural inferenee is that the Walls of tllO latUT coalesce with tin' walls of the sinuses to form this intervening mem- brane. In this manner the embryonic and maternal structures are inseparably Mended. ! 1 . I-,. assu with the growth of the embryo, while the womb expands in concert with this action in the chorion and embryo. UNIVERSITY OF CALIFORNIA LIBRARY, BERKELEY THIS BOOK IS DUE ON THE LAST DATE STAMPED BELOW Books not returned on time are subject to a fine of 50c per volume after the third day overdue, increasing to $1.00 per volume after the sixth day. Books not in demand may be renewed if application is made before expiration of loan period. SEP 16 1930 NOV 2 4 1939 & 60m-7,'29 jrtg 399398 UNIVERSITY OF CALIFORNIA LIBRARY