Introduction Chapter 1 : The reproductive organs and the formation of the sex cells oviduct ampulla uterus ovarium omentum latum Fallopian Funnel cervix vulva vagina The entire system lies in the pelvic region of the abdominal cavity The uterus is thick walled and consist of two layers the inner endometrium and the outer muscular myometrium The embryo imbeds in the endometrium by means of the placenta The ligament, the omentum latum hold all the different structures together The ovarium consists of a germinal epithelium surrounding the connective tissue the stroma The vagina acts as the copulatory organ and the birth canal During pregnancy, the sphincter muscles of the cervix remains contracted. The remaining canal is blocked by means of a gelatinous plug The embryo implants in the uterine wall and connection with the mother is established by the placenta (foetal & maternal) tissue vagina •placenta performs all the necessary functions; •provides nutrition; •supplies oxygen and •removes carbondioxide; •removes waste products Position of the testes is outside the body cavity The spermatozoa (sperm) die off at normal body temperature. Ligaments are relaxed to hang the testes away from the body but will contract at temperatures below the optimal, to bring the testes close to the body. Exceptions are the elephant and the springhare Attached to each testis is the epididymis, which is expanded into the deferent duct The two deferent ducts enter the body cavity, loop around the bladder and fuse together and with the urethra, while passing through the following glands: Seminal vesicles (2x) Prostrate gland (1 consisting of several lobes) Cowper’s gland (= bulbo-urethral gland) Secretions of the glands are added during ejaculation to produce the semen Functions of the secretions are: Maintain correct pH Provide volume/liquid medium Provide nutrients The urethra extends trough the penis Testis is divided into compartments by septula testis Each compartment is packed with seminiferous tubuli and interstitial cells Interstitial cells secrete testosterone – carried through the bloodstream Responsible for development of male characteristics Immature sex cells develop in the walls of seminiferous tubuli Mature sperm come free; move along efferent ducts into the epididymis (caput, corpus & cauda epididymis) Efferent ducts fuse to form deferent duct that leaves epididymis and enters the body cavity Interstitial cells secrete testosterone – carried through the bloodstream Responsible for development of male characteristics Immature sex cells develop in the walls of seminiferous tubuli Mature sperm come free; move along efferent ducts = process of the formation and development of gametes Number of chromosomes are halved during meiosis to form spermatozoa and ova Spermatozoa and ova are haploid The development of the male gamete is spermatogenesis The development of the female gamete is oogenesis 1 Male primordial generative cell (spermatogonium) results in 4 spermatozoa 1 Female primordial generative cell (oogonium) results in one ovum + 3 polar bodies New spermatogonia are formed throughout the life of the male from puberty on Spermatogenesis commences at puberty and continues into old age All the stages of spermatogenesis are represented in the seminiferous tubuli at any given time The maturation process from spermatogonium to spermatozoon takes about two months Spermatogonia lie dormant in seminiferous tubuli against basement membrane Continual mitotic division since puberty Some cells start growth process – 1 primary spermatocyte Meiosis I – 2x secondary spermatocytes = ½ size of primary spermatocyte Meiosis II – 4x spermatids = ½ size of secondary spermatocyte Spermatogonia lie dormant in seminiferous tubuli against basement membrane The cells move away from basement membrane while growth and meiosis progress & lie among Sertoli cells Spermatids lie against the lumen of seminiferous tubule Process of differentiation follows Normal spherical spermatid becomes a free-swimming sperm Granules become visible in Golgi body Vacuole + granules lie close to nucleus Vacuole of Golgi body enlarges Granule becomes visible and enlarges to form the proacrosomal body Liquid moves away and form the Golgi rest Membrane of vacuole spreads out over nucleus = cap Nucleoplasm with organelles flows to one side Just a thin layer of cytoplasm remain covering the spermatozoon Golgi Rest + excess cytoplasm is discarded Pronucleus + acrosonal granule = head of the sperm Pronucleus elongates Spermatid elongates Centrosome contains 2 centrioles Centrioles move to opposite side of pronucleus Become proximal & distal centrioles Proximal centriole lies in depression in nucleus (forms 1 of the asters during 1st division) Axial filament grows out of distal centriole (forms the basal granule) Axial filament consist of 1 pair of central & 9 pairs of fibres situated around the circumference Mitochondrion forms a spiral around the axial filament (provides energy) At the end of the mitochondria is the ring centriole (function unknown) From proximal centriole to ring centriole = middle piece Axial filament elongates form flagellum Flagellum is covered by a thin layer of cytoplasm and a sheath in mammals The sheath extends from the ring centriole to almost the tip of flagellum The tip of the flagellum is naked. Sheath consists of 9 fibres twisted around flagellum Process start before birth of baby girl = Prenatal maturation Cells of germinal epithelium divide through mitosis to form oogonia Some grow to form primary oocytes that separate from epithelium and move into stroma Cells from the stroma known as follicle cells surround each of the primary oocytes to form a number of primordial follicles Meiosis I starts but process is arrested in Prophase I before birth of girl baby, due to the secretion of a hormone, Oocyte Maturation Inhibitor (OMI) by the surrounding follicle cells The girl baby is born with all the primary follicles she will ever have All the primary follicles remain suspended in Prophase I until puberty, when Postnatal maturation commences. Process is stimulated by Follicle Stimulating Hormone (FSH) secreted by the pituitary gland at the onset of puberty The pituitary is situated ventral and attached to the brain by a stalk From puberty on, one primary follicle will enlarge and move deeper into the stroma, approximately once every 28 days in one of the ovaria The primary oocyte now proceeds with Meiosis I while the follicle develops Follicles in different stages of maturation can be found in the ovaries of the reproductive female at any time The follicle cells around the primary oocyte become collumnar and start secreting estrogen = primary follicle The influence of estrogen is: (i) development of female characteristics & (ii) build up of uterine lining The follicle layer becomes multi-layered = secondary follicle An eccentric cavity, the follicular antrum, filled with fluid probably secreted by the follicle cells, appears within the follicle cells = Graaffian/ tertiary follicle The young Graaffian follicle moves further into the stroma, and the stroma lays down a tough capsule, the theca folliculi around it The young Graaffian follicle grows as more fluid collects in the antrum, pushing the dividing primary oocyte surrounded by some follicle cells against the side of the Graaffian follicle Gradually the follicle moves back towards the surface of the ovarium A membrane, the zona pellucida is secreted by the follicle cells around the dividing primary oocyte Meiosis I is completed just before the Graaffian follicle opens to release the secondary oocyte surrounded by the zona pellucida + follicle cells into the body cavity = ovulation The division of the cytoplasm is unequal. One daughter cell receives most of the cytoplasm = secondary oocyte. The other receives mostly nuclear material = polar body Polar body is non-functional & might divide again (2 polar bodies) before degenerating Ovulation is regulated by 2 hormones secreted by the pituitary, FSH and the luteinizing hormone (LH) Meiosis II commences just after ovulation, but stops in metaphase II. It is only completed after a sperm has penetrated the cell membrane of the secondary oocyte The Meiosis II division also results in a cell that retains most of the cytoplasm = ovum + a polar body The ovum that contains a large quantity of cytoplasm is larger than any other somatic cell in the body Oogenesis is completed by the completion of Meiosis II, while the sperm is already contained in the cytoplasm of the ovum. The follicle cells of the ruptured Graaffian follicle secretes a large quantity of estrogen into the bloodstream. The blood supply to the uterine wall is increased under the influence of the large quantity of estrogen The ruptured Graaffian follicle proceeds to grow, becoming a temporary endocrine gland, the corpus lutheum The corpus luteum remains throughout pregnancy, secreting the hormone progesterone. Progesterone inhibits ovulation during pregnancy and for sometime thereafter No pregnancy – the corpus luteum will degenerate after about 10-12 days = corpus albicans Towards the end of pregnancy, the secretion of progesterone decreases, a birth process commences = corpus albicans No of primary oocytes at birth of female infant = about 2 million 30 – 40,000 at puberty Only approx 400 become secondary oocytes and are released by the ovaria Female reaches menopause (end of reproductive life) after last ovulation Age – 45 to 50 years The ovum (>somatic cells) contains haploid no of chromosomes (pronucleus) + usual cytoplasmic organelles In mammals it is surrounded by the zona pellucida In addition it containes: Superficially situated cortical granules Yolk only in animals that lay eggs – absent in mammals Brown or black pigment granules NB. The size of the ovum is relative to the amount of yolk it carries i.e. Bird/reptile egg is large compared to the small mammalian egg. 1. Primary egg membranes = membranes formed by the female cell 1.1 Plasmalemma = membrane of female sex cell 1.2 Vitteline membrane – found in most animals; absent in mammals = thin & transparent lying close to the plasmalemma, Forms the fertilization membrane during the zona reaction, where it is present after penetration. 2. Secondary egg membranes = membranes secreted by the follicle cells (only found in mammals) 2.1 Zona pellucida – clear, thick membrane secreted by the follicle cells. Transformed into the fertilisation membrane during the zona reaction 2.2 Corona radiata – consists of the follicle cells that surround the secondary oocyte after ovulation. Follicle cells are held together through hyaluronic acid. These cells gradualy disperse after fertilization and before implantation 3. Tertiary egg membranes = secreted by the oviduct/glands of the oviduct. Absent in placental mammals 3.1 Jelly – around amphibian eggs 3.2 albumin & leathery or hard shells of reptile and birds Chapter 2 : Fertilization Stages: 1. Fertilisation 2. Cleavage 3. Gastrulation 4. Neurulation & Formation of the primary organ rudiments 5. Organogenesis 6. Growth Ovulation places sec oocyte in the body cavity. The movement of the fimbriae of the Fallopian funnel creates a current in the peritoneal fluid in the body cavity and the sec oocyte is swept into the infundibulum of the oviduct. The epithelial cells of the oviduct are ciliated, and these cilia move the sec oocyte into the ampulla part of the oviduct About 200 – 500 million sperm is deposited in the area around the opening of the cervix The sperm is stored in the mucus and crevices of the cervix & released over a period of 3 days The liquid contents of the vagina breaks down the plasmalemma of the head of the sperm = capacitation (about 7 hours) The acrosomal granules are now released, become dissolved, forming enzymes called sperm lysins Sperm lysins have the ability to dissolve the egg membranes In some animals a long rigid filament form (centre of the acrosome) known as the acrosomal filament Acrosomal filament is absent in man – plasmalemma becomes perforated to release the spem lysins While capacitation takes place the sperm is carried through the cervix, lumen of the uterus and up the oviducts to the ampulla on both sides The sperm collide with the sec oocyte by chance: • Sperm can move, but the direction is random. •Many sperm are present •The egg is very large Large quantity of sperm reach the ampulla. The sperm become ‘sticky’ – adheres to the corona radiata of the sec oocyte & to each other. Excess sperm is neutralized in this way Corona radiata releases fertilizin & surface cytoplasm of sperm releases antifertilizin •Fertilizin + antifertilizin = chemical bond •Fertilizin = glycoprotein/mucopolysaccharide •Antifertilizin = small acid protein •Fertilizin & antifertilizin is species specific i.e. will only combine with the other of its own species Chemical process Hyaluronic acid (mucopolysaccharide) hold follicle cells together = corona radiata Perforated acrosome of agglutinated sperm secretes enzyme hyaluronidase Hyaluronidase dissolves hyaluronic acid & follicle cells falls apart to allow sperm to reach zona pellucida Perforated acrosome of agglutinated sperm secretes enzyme acrocin that dissolves zona pellucida – sperm head reaches plasmalemma Plasmalemma of sperm head & plasmalemma of sec oocyte fuse, breakdown – contact between cytoplasm of gametes Head of sperm = male pronucleus + flagellum swims free, while rest of sperm i.e. plasmalemma, sheath of tail & thin layer of cytoplam remain attached to plasmalemma of sec oocyte Secondary oocyte completes meiosis II Flagellum of sperm degenerates and disappears Head of the sperm recovers fluid from the female cytoplasm – male pronucleus Male pronucleus rotates 180 degrees and approaches with prox centriole Nuclear membranes break down to form the zygote (2N) Chromosomes become visible on equator = 1st mitotic division = cleavage Triggered by the passage of the sperm through the zona pellucida and starts at the point of penetration, gradually spreading over the surface of the sec oocyte Cortical granules take fluid from the cytoplasm and explode. Membrane of the cortical granule ruptures to release its contents = lysosomal enzymes into the space between the plasmalemma and the zona pellucida. Space created is perivitteline space Lysosomal enzymes act on zona pellucida – becomes impereable to other sperm = fertilization membrane Process takes 20 minutes Second temporary change involves a change in electrical potential of the inside surface of the plasmalemma The electrical charge is normally – Upon penetration the electrical charge changes to + The electrical potential of the sperm head is + I.e. any other sperm head will be repelled by the ovum plasmalemma The process takes 2 seconds The ovum plasmalemma’s electrical charge changes back to – as soon as the fertilization membrane is in place The receptivity of the egg to penetration is reduced to 1/120 by the zona reaction 1. Monospermia – 1 sperm penetrates the sec oocyte in most groups due to zona reaction 2. Pathological polyspermia – 2 sperms penetrate due to high density of sperm – triploid zygote that cannot divide and dies off 3. Physiological polyspermia – Found in birds. It is normal to find more than one sperm in the cytoplasm of the ovum, but only 1 male pronucleus fuse with the female pronucleus, while the others die off. Chapter 3 : Morphogenesis or Ontogeny Stages: 1. Fertilisation 2. Cleavage 3. Gastrulation 4. Neurulation & Formation of the primary organ rudiments 5. Organogenesis 6. Growth • • • • • • Single celled zygote is transformed into a multicellular body through rapid mitotic divisions The characteristics of the cleavage divisions are: No growth occurs Shape remains the same (sphere = blastula) A cavity originates inside the blastula = blastocoele No qualitative changes in the cytoplasm just transformation of food reserves into active cytoplasm & cytoplasmic substances into nuclear substances Posistion of parts of cytoplasm remains in the same positions Ratio nucleus:cytoplasm is normalised Different cleavage types are determined by: 1. Presence and amount of yolk in the cytoplasm 2. The phylogenetic position of the group Eg. Branchiostoma / the lancelet / Amphioxus The zygote contains very little yolk Divisions are complete Daughter cells are of equal size 1st division is vertical – 2 identical blastomeres 2nd division is vertical - 4 identical blastomeres 3rd division is horizontal – 8 blastomeres; 4 micromeres & 4 macromeres 4th division is vertical – 8 micromeres & 8 macromeres = 16 cell stage 5th division is horizontal – 32 cells = morula Further divisions – blastula: blastoderm surrounding a blastocoele Blastoderm is a single layer of cells Thinner in the animal hemisphere than in the vegetal hemisphere Eg the amphibian egg The zygote contains a moderate amount of yolk Divisions are complete, but it takes time to complete the division The initial four daughter cells are similar in size, but daughter cells of following division differ in size The amount of yolk included in the cells slow down the division processes 1st division is vertical 2nd division is vertical 3rd division is horizontal Blastula consists of a multilayered blastoderm and an eccentric blastocoele in the animal hemisphere Blastoderm cells in the vegetal hemisphere are larger and fewer that the cells in the animal hemisphere Eg bony fish, birds and reptiles The egg contains a large amount of yolk Divisions are incomplete i.e. the cell membrane that separates the two daughter cells are never completed The incomplete cleavage divisions only takes place in the animal pole where the yolk load is very heavy to form a blastodisc Eg. placental mammals The egg contains no yolk Cleavage is a series of rapid, complete, non–synchronised mitotic divisions resulting in progressively smaller blastomeres i.e. stages like 5 and 7-cell stages can be seen Cleavage results in a blastocyst The blastocyst consists of a trophoblast, a blastocyst cavity and some formative cells, called the inner cell mass Stages: 1. Fertilisation 2. Cleavage 3. Gastrulation 4. Neurulation & Formation of the primary organ rudiments 5. Organogenesis 6. Growth Gastrulation is the process whereby the blastula/ blastodisc/ blastocyst is transformed into the three germ layers; the ectodem; mesoderm; endoderm All the tissues of all chordate animals originate in these three germ layers: Ectoderm – neural tissue i.e. nervous system and epidermis i.e. skin Mesoderm – support system i.e. (skeleton and muscles; cardiovascular system (heart, blood vessel & blood; urogenital system (kidneys & reproductive systems) Endoderm – alimentary canal and associated organs (lungs, liver, pancreas) The process of gastrulation like cleavage differs between the different classes of the vertebrates again according to: The amount of yolk contained in the cytoplasm of the zygote and The phylogenetic position of the class Chapter 4 : The Development of Branchiostoma Cleavage = Equal holoblastic cleavage The future of the blastoderm can be plotted The animal hemisphere cells are mostly yolk free and is destined to develop into ectoderm Most of the vegetal hemisphere cells are contain yolk, the cytoplasm is granular and is destined to develop into endoderm A small crescent of loosely packed cells in the vegetal hemisphere is also yolkfree like the future ectodermal cells, but stains with basic stains. These cells are destined to become mesoderm which includes the future notochord material The single walled blastula needs to become an embryo consisting of 3 layers Gastrulation takes place through the invagination of the future endodermal and mesodermal cells The 1st sign of gastrulation is the flattening of the vegetal pole (endoderm) The endoderm invaginates into the blastocoele followed by the mesoderm The mesoderm includes both the future muscle tissue and the rudiment of the notochord The hollow sphere now becomes a double walled cup; the meso & endoderm forming the inner layer, and the ectoderm situated in the outer layer •The continuation of invagination obliterates the blastocoele and a new cavity the archenteron is formed. •The embryo is now called a gastrula •The rim of the double walled cup is at first wide, but it contracts to form the blastopore, enclosing the endoderm and the mesoderm. •The mesoderm includes the muscle tissue and the rudiment of the notochord •The notochordal tissue converge on the mid-dorsal line with the remaining mesodermal tissue on either side •A pear-shaped part of the ectoderm on the mid-dorsal line flattens under the influence of the underlying notochord to become the neural plate. The appearance of the neural plate shows that the process of neurulation has started. The neural plate separates from the surrounding ectoderm, sinks in and becomes covered by the remaining ectoderm, now called the epiderm. •The epiderm will become the integument or skin •The archenteron is lined dorsally by the notochord that forms a longitudinal band on the mid-dorsal line with the mesodermal tissue on either side. •The lateral and ventral walls of the archenteron are formed by the endoderm • The presumptive tissues of the different germ layers separate from each other through the formation of crevices • The notochord band becomes a solid rod • The future muscle tissue separates from the endoderm through the formation of a groove in contact with the archenteron. This groove closes off from the archenteron and the tube thus formed breaks up into segments • Each of these segments surrounds a cavity and is called somites and the process is metamerisation. The cavities in the somites will become the coloem •The neural plate rolls itself into a tube, the neural tube. •All the primary organ rudiments have been laid dowm. •From here on the body of the lancelet will elongate further, the blastopore becomes the anus, while the mouth breaks through just anterior to the tip of the notochord. Chapter 5 : Early development of the avain embryo and the extra-embryonic membranes Cleavage is typical Meroblastic, discoidal or Incomplete The egg contains a large amount of yolk Divisions take place in the animal pole and are restricted to the cytoplasmic cap where a little of cytoplasm surrounds the nucleus of the zygote Divisions are incomplete i.e. the cell membrane that separates the two daughter cells are never completed The first divisions are all vertical, the daughter cells are separated by the cleavage furrows but remain open to the yolk Later on horizontal divisions take place and a slit-like cavity (comparable with the blastocoele)is formed between the blastomeres and the yolk to form the blasotodisc • The entire embryo develops out of the blastodisc. As nutrients re accumulated and need to be provided, waste products needs to be stored, gaseous exchange need to be facilitated and the embryo need to be protected against desiccation and mechanical shock extra-embryonic membranes also develop out of the blastodisc •There are 4 extraembryonic membranes: 1. The amnion – envelopes the embryo and filled with amniotic fluid 2. The yolk sac that envelops the yolk with fingerlike pojections that penetrate deep into the yolk 3. The allantois - an outgrowth of the hind-gut that spreads out between the amnion, yolk sac en the overall enveloping chorion (4) • The amnion, allantois and yolk sac is connected to the embryo by means of the umbilical chord • Embryonic bloodvessels run through the umbilical chord to the amnion, yolk sac and allantois 1. Amnion – (i) prevents desiccation and (ii) protects the embryo against mechanical shock 2. Yolk sac – (i) Absorbs the yolk into the bloodstream and (ii) Transports the nutrients to the embryo 3. Allantois – (i) stores the waste products inside the allantoic cavity and (ii) absorbs the oxygen from the air that enters through the shell, shell membranes and chorion and stransports it to the embryo and transports the CO2 from the embryo away from the embryo so that is can move out 4. Chorion – (i) protects the embryo and the other extraembryonic membranes from friction against the hard shell and (ii) from adhesion to the shell Chapter 6 : The development of the mammalian embryo •Contains NO yolk •Embryo is retained in the uterus •Attached to the wall of the uterus by means of a new structure the placenta •The placenta facilitates gaseous exchange, provides nutrition and removes waste products • Fertilization takes place in the ampulla of the oviduct • Cleavage takes place as the zygote moves down the oviduct towards the uterus •The morula enteres the uterus 3 days after fertilization •The morula is 12 – 16 cells forming a solid ball of cells and is still surrounded by the zona pellucida • As cleavage divisions continue cavities appear in the interior of the morula The cavities become filled with fluid from the uterine cavity to become the blastocyst cavity The embryo is now a blastocyst, consisting of the trophoblast, surrounding the blastocyst cavity with the inner cell mass/embryoblast/ formative cells at the embryonic pole The inner cell mass is basophilic-stains bright when stained with basic dyes. The trophoblast gives rise to the extra-embryonic membranes, while the embryo proper develops out of the inner cell mass. The zona pellucida (fertilization membrane) is still intact around the blastocyst and the blastocyst lies free in the uterine cavity. The zona pellucida starts to degenerate and eventually disappears. This enables the blastocyst to become attached to the endometrial epithelium. The embryo is now referred to as a conceptus. The conceptus orientates itself in such a way that the embryonic pole lies against the endometrium of the uterus. The trophoblast attaches itself to the endometrial epithelium in the following way. The trophoblast cells in contact with the endometrial epithelium undergo multiple divisions Differentiates into two layers: inner cellular layer = cytotrophoblast and an outer = syncytotrophoblast. Cell membranes of the outer layer disappear to form a multi-nucleated protoplasmic mass = syncytium Forms finger-like processes which grow into the endometrium. The embryo is superficially implanted by the end of the first week. • A new layer becomes visible during attachment and superficial implantation • A flat layer of cells possibly originating from the trophoblast arranges, through a process of migration, into a thin layer of flat cells lining the inner surface of the inner cell mass to form the hypoblast. • The hypoblast therefore forms a ceiling to the blastocyst cavity, separating the inner cell mass from the blastocyst cavity. The inner cell mass + hypoblast arrange into a double layered disc = the embryonic disc, consisting of a top layer of columnar cells = the epiblast and a second layer of cuboidal cells, the hypoblast. Also known as the bilaminar embryonic disc. The epiblast develops into all three germ layers of the embryoproper. The cells of the hypoblast develop into endoderm or part of it. It is believed that its contribution is mostly extra embryonic in that it mainly contributes to the endoderm of the extra embryonic membranes. The syncytotrophoblast invades deeper into the endometrial stroma. The cells of the stroma degenerate in the area of the penetrating syncytotrophoblast. The trophoblast continue to divide producing cells that migrate into the increasing mass of the syncytotrophoblast where they soon loose their cell membranes. The formation of the syncyto trophoblast expands around the cytotrophoblast of the conceptus as implantation progresses. Cavities form between the inner cell mass and the trophoblast. These spaces join each other to form the amniotic cavity. The part of the cytotrophoblast overlying the epiblast splits into 2 layers. The inner layer consists of amnioblast cells The amnioblasts & cytotrophoblast together becomes the the amnion. The cells of the epiblast form the floor of the amniotic cavity and are continuous peripherally with the amnion. The cells of the hypoblast form the roof of the blastocyst cavity and are peripherally continuous with the cytotrophoblast. Delamination of the cytotrophoblast surrounding the blastocyst cavityalso takes place The inner layer of the cytotrophoblast around the blastocyst cavity is continuous with the hypoblast It is a thin membranous layer called the exocoelomic membrane The blastocyst cavity is now referred to as the exocoelomic cavity. Cavities appear in the syncytotrophoblast called lacunae which soon become filled with maternal blood from the ruptured capillaries and secretions from the eroded endometrial glands. This nutritive fluid is known as embryotroph and reaches the embryonic disc by diffusion. The lacunae become connected with the maternal blood vessels and in this way the uteroplacental circulation is established. Oxygenated blood reach the lacunae through the arteries, and the deoxygenated blood is removed from the lacunae by the uterine veins, creating a primitive uteroplacental circulation. A lacunar network is established giving the syncytotrophoblast a sponge-like appearance. The lacunar network is initially formed in the area of the embryonic pole. This network increases and spreads throughout the whole endometrium to surround the conceptus completely. At this stage cavities appear in the syncytotrophoblast called lacunae which soon become filled with maternal blood from the ruptured capillaries and secretions from the eroded endometrial glands. This nutritive fluid is known as embryotroph and reaches the embryonic disc by diffusion. The lacunae become connected with the maternal blood vessels and in this way the uteroplacental circulation is established. Oxygenated blood reach the lacunae through The conceptus continues to penetrate deeper into the stroma of the endometrium until it is completely embedded. A closing plug consisting of a blood clot and cellular debris remain on the surface of the epithelium for a short while until the regenerating epithelium covers over the conceptus. This type of implantation where the entire conceptus is embedded into the endometrium of the uterus is called interstitial implantation.