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    10 views42 pages

    Circulatory System Pattens

    Uploaded by

    Zahra Nur Azizah

    Copyright:

    © All Rights Reserved
    Download as PDF or read online from Scribd
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    of 42
    spe CHAPTER f 8 eee : THE DEVELOPMENT OF THE CIRCULATORY SYSTEM INTRODUCTION TO THE EMBRYONIC CIRCULATION ‘The plan of the embryonic heart and vascular system‘is bound by three major constraints. First it must mect the immediate needs of the embryo at its various stages of development by supplying it with oxygen, nutrients, and other essential materials for growth while at the same time removing CO and other metabolic wastes. For this, only a simple unidirectional pumping action is required of the heart to move the blood along channels supplying the develop- ing organs of the embryo. To meet the requirements of respiration, nutrition, and excretion, two extraembryonic circulatory arcs have developed (Fig. 8-17). The arc to the yolk sac, the vitelline arc, supplies foodstuffs to all large-yolked vertebrate em bryos; however, in mammals the yolk sac and vitelline circulation persist for what were originally probably subsidiary functions, the origin and transport of primordial germ cells and primitive blood cells. The large allantois of amniote eggs (with its circulatory arc) was originally the chief organ of respiration and deposition of excretory wastes, but as the placenta in mammals evolved into a more efficient organ of exchange, the allantois correspondingly became re duced in prominence. The allantoic circulation, however, has been incorpo rated into the placenta and continues to serve its original functions. In addition to satisfying the relatively simple requirements of the embryo the plan of the embryonic circulation in amniotes must also anticipate the ee needs of the embryo once it hatches from the egg or is delivers’ « mother's uterus (Fig. 18-1). The most immediate and critical adjust” ment to birth is the need of the new! aaa F tly. i 4 ly born dependently: Breathing requires that the lunge 2 re ae aoe to breathe indep: ¢ developed tardily from the 586 Dipindai dengan CamScanner rte San Risht ventricle, Frlnonery tp by, a, rel ie | 5 ES set » track, hee Eh Pete Ureter, ebete tinker Pas Ades Teles Serie FIGURE 16-1 i Pian of tne postnatal circulation. (Aer Paton, 1963, nFsoan Bit ete eae nal Foundation and the J. B. Lippincott Company. Th ee eat eng UGE ere important fetal vessels (Fe 1420) wn a ” acu reduced to fibrous cords. indicates the vatwla foram vats ne soca poston garda foc posal ua (See cored set) 587 Dipindai dengan CamScanner 588 THE DEVELOPMENT OF THE CIRCULATORY SYSTEM morphological standpoint and are untested functionally, begin to function at full capacity immediately after birth. Because of this, the embryonic heart cannot be content with remaining in its original condition, as a simple tube with the blood passing through it in an undivided stream, Early in embryonic life it must be converted into an elaborately valved four-chamber organ, partitioned in the midline and pumping from: its right side a pulmonary stream which is retumed to the left side and pumped out again as a systemic bloodstream, Ang the heart cannot cease work while making its internal alterations; there can be no interruption in the current of blood which it pumps to the growing embryo, The systemic, as well as the pulmonary, part of the circulation must be Prepared. Because of the delayed development and restricted vascular bed of the embryonic lungs, the left side of the heart receives less blood from the Pulmonary veins than the right side of the heart receives from the venae cavae, Yet after birth the left ventricle is destined to do more work than the right ventricle. On the other side of the coin, the right ventricle receives More blood than can be accommodated by the pulmonary circulation, These and other problems of the embryonic heart are solved by the presence of shunts, which act like safety valves, allowing the various chambers of the heart to obtain the exercise they need for their Tequired development but not overloading the Pulmonary vasculature beyond its limited carrying capacity, Besides meeting immediate and future physiological needs, the circulatory Dipindai dengan CamScanner EMBRYONIC HEMATOPOIESIS 599 FIGURE 18-2 Sites of primary : erythropoiesis (shaded gray) in amphibian (A) and bird (B) ‘embryos. again to functional significance, for the relationship of the aortic arches to the sill arches writes into the story of individual development an unequivocal record of the evolutionary phase when the gills were a center of primary metabolic importance. Other examples of recapitulation in the circulatory system are the previously mentioned persistence of the vitelline circulatory arc long after the yolk sac has abandoned its original nutritive function and the series of highly developed venous channels in the mesonephros, even when it is destined to degenerate later in development. EMBRYONIC HEMATOPOIESIS The first blood cells are produced in extraembryonic sites as small groups of mesodermal cells called blood islands (Fig. 18-2). The early ood islands are located next to the endodermal wall of the yolk sac (Fig. 18-3), and in : chic! the differentiation of blood islands has been shown io depend on af ericron between the splanchnic mesoderm and the underlying en’orr , 1965). alan island become flattened-as a Cells in the outer zone of the primordial Seed eeeay located cells, Young vascular endothelium and SPEC ig.3B). Within the endothelial which become hematopoietic stem cells (Fi IO). Nery Cay! Vesicles, fluid accumulates and suspends the de\ ne onoecopoiess in the There is a temporal progression of maior se" Ce Cen ig the yolk s3c. el .-6). The first hematopoietic Het" yund in the liver and aes a dominant sites of hematopoiesis re eet experiments spleen, body mesenchyme, and Sinai te Oh toa chick yolk sae) have shown oie bij sail body 8F etic sursor cells in the on chimeric bird embryos Ke ulations of ‘hematopoietic breenibryonic) arises ae eng Se ra mbryo (Dieterien-Lievre. ! Dipindai dengan CamScanner 590 THE DEVELOPMENT OF THE CIRCULATORY SYSTEM & 4 Unb. ven veusanan spt ani abet FIGURE 18-3 MINP Bog Yolk. sepment ot yok-sac blood islands in human embryus. A-C are camera lucida drawings, re coders eae (A) Early stage in aggregation of cols behvncy endoderm and splanchrie me: in yok sae ofan embryo early in fourth week {17 somites). (B) Beginning of diferenta- fon ol endothe and primitive blood cells rome embryo of about 4 weeks (4.5 mm). (C) A cles euaponeey, net om a 4:Week embryo showing ‘endothelium well differentiated and corpus: Hee FeSBENGEd, feo, in plasma. (0) The Doone somite embryo showing location of young blood islands on yok sac, (Camegie Cont. to Emb., 1829, vor 20) Abbreviation: Heb., primitive blood-mother-cet, or hemocytoblent in the yolk sac and constitut new population of hémato mesoderm, produces a f erythrocytes that Progressively replaces the yolk sac-derived Sothrocytes. The cells in the liver and spleen, and ultimately the bone Marrow, are those which give rise to the erythroid series (Cells forming red blood cells, or erythrocytes) and the granulocytic series (cells lated White blood Cells, or neutrophils, eosinophils, and ntral lym ins, the thymus and the bursa of Fabri= id cells (lymphocytes and monocytes): : I iia 8 in the central lymphoid organs, In later life other pa eed the peripheral lymphoid tissues (see Chap. 19. Dipindai dengan CamScanner ERYTHROPOIESIS ANO HEMOGLOBIN FORMATION 591. envTHROPOIESIS AND HEMOGLOBIN FORMATION Enthropoiesis is the process by Which a mature red blood cell (erythrocyte) faaded with hemoglobin molecules differentiates from primitive hemocyto- Hastie stem ceils. Erythropoiesis can be viewed from several aspects and jevels of organization. At the tisste and organ level, there are three major phases in erythropoiesis (Fig. 18-6, bottom). For a brief period the only Hood-forming activity occurs extraembryonically, in the yolk sac. After the cecond month the major sites of erythropoiesis shift from the yolk sac to jintraembryonic organs. The second major phase is the hepatic period {roughly the third through seventh months of human pregnancy), during which the liver and spleen are the dominant hematopoietic organs. Finally, late in pregnancy the bone marrow takes over as the defi erythropoiesis in higher vertebrates The differentiation of individual erythrocytes must also be viewed in the context of their site of origin. The first erythrocytes are derived from the original population of yolk-sac precursors. These cells differentiate relatively synchronously and are released into the bloodstream at an early stage of differentiation. Maturation is completed in the bloodstream, and in mammalian embryos the yolk sac-derived erythrocytes are nucleated. Differentation of the erythrocytes derived from intraembryonic precursor cells has been extensively studied with respect to both cellular morphology and production of hemoglobin. Both light and electron microscopic prepara~ tions show a series of developmental stages which are classical for a cell that is heavily engaged in the production of intracellular protein (Fig. 18-4). From the hemocytoblast, a primitive cell that is considered by many.to be the stem cell for all families of blood cells, the erythrocyte line first appears as a highly basophilic cell called a proerythroblast. These cells have undergone sufficient restriction to be firmly committed to the production of red blood cells, but they have not yet begun to produce hemoglobin in amounts sufficient to a detected by cytochemical analysis. They have large nuclei with prominet Aucleoli and largely uncondensed nuclear chromatin. Synthesis of globin TRNA is high. The cytoplasm contains aggregates of mainly free ribosomes, Which will be used in intracellular protein synthesis. hromato- Subsequent stages of erythroid differentiation (basophilic, polychromale. Philic, and orthochromatic erythroblasts) show a progressive change 1 (it balance between the accumulation of newly synthesized hemoglobin Ms Sules and the decline of first the RNA-producing pa een continu aejeit-syathesizing apparatus. During ieee ane (eharacteristic of. usly stains less basophilically as its eosinophil i (oars changes Sccumulated protein) increases in intensity. Corresponding Morr ¥S decrease in the concentration of ribosomes in the cytoplasm tay Shows signs of its inexorable pathway toward inactivation ac tary ig its steadily decreasing size, the progressive condensation OT atic and the elimination of the nucleolus. Finally, the late Ik-sac erythro- Srythroblast extrudes its pycnotic nucleus. In contrast to yol ive site for Dipindai dengan CamScanner svsTeM yt oF THE CIRCULATORY 592 THE DEVELOPME! Proerythroblas, ile Polychromatopti erythroblast Orthochromatie exythroblast Erythrocyte Reticulocyte FIGURE 18-4 : i atic i i hemogle- Suocessive stages in the diferentiation of an erythrocyte. increasing concentrations of bin are indicated by the Intensity of the gray shading in the cytoplasm. {Adapted from Rifkind, 1874 in Lash and Whittaker, eds,, Concepts of Povelopment, Sinauer Assoc.) * Poiesis, the changes normally occur within the hematoy Only after it has lost its nucleus is the cell, into the bloodstream, Reticulocytes sti Somes, and they Continue poietic tissues, and ustice to the extent to which the ed to fulfilling its function. 7 Dipindai dengan CamScanner ERYTHROPOIESIS AN HEMOGLOBIN FORMATIgN 593 Chromosome 11 Y fore = i ( 2 om Major aut FIGURE 10-5 Seauenial geno activation along chromosomes 11 and 16 In he- denahin synthesis curing development. (Ater &. Gibor, 108s Developmental Biology, Sinaver) There are developmental changes in the hemoglobin molecule itself. The hemoglobin molecule (MW 64,500) contains four polypeptide chains (giobin) ChuPlexed to a molecule of heme. The human globin genes are located on fhromesomes 11 and 16 (Fig. 18-5), and during embryonic development there Gnu oderly succession of pairs of globin types (with a pair derived from each {hfomosome) contributing to the hemoglobin molecule. The earliest hemoglo- bin molecule (embryonic hemoglobin) contains a pair of {globin molecules derived from chromosome 16 and a pair of eglobin chains from chromosome 11 jecched to the heme molecule. A pair of a-globin molecules, specified by genes located on chromosome 16, soon supplants the original pair of — ati ane from this point on, regacaless of the type of hemoglobin, two of the globit Shains are always a molecules. 7 asEbtvonic hemoglobin is present for only a couple of months, after which

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