(ii) It helps to retain sufficient amount of cytoplasm in the ovum which is essential for the development of early embryo
(iii) During meisosis first crossing over takes place which brings about variation
end is vegetal pole. Cytoplasm of ovum is called ooplasm. It has a large nucleur or germinal vesicle.
TYPES OF OVUM
On the basis of amount of yolk, eggs are classified as:
(i) Alecithal – No yolk, example : human egg
(ii) Microlecithal – Small amount of yolk example : Sea urchin
(iii) Mesolecithal – Moderate amount of yolk, example frog and other amphibian egg
(iv) Macrolecithal or polylecithal – Large amount of yolk example : reptilian and avian eggs
• On the basis of distribution of yolk, eggs are classified as:-
(i) Isolecithal or Homolecithal _ Having homogenously distributed yolk, example : Protochordates and echinoderms.
(ii) Hetrolecithal – egg with unevenly distributed yolk
(iii) Telolecithal – Having yolk concerned in one half Example : amphibian eggs
(iv) Centrolecithal – yolk is concentrated in centre and cytoplasm is peripheral example – insect eggs
(v) Discoidal or Meiolecithal – Almost the whole of the egg is occupied by the yolk except a small disc, example : eggs of birds and reptiles
• On the basis of presence and absence of shell eggs are differentiated into cleidoic ( surrounded with water proof shell e.g. birds and reptiles) and noncleidoic ( shell absent)
FERTILIZATION IN HUMANS
• It is fusion of male and female gametes to form zygote. In human being fertilisation is internal. Human beings are viviparous
• Here the embryo is retained and nourished inside the uterus of the female by means of an attachment called placenta. At one time only a single ovum is released in human females from one of the two ovaries towards the middle of ovarian/ menstrual cycle. It passes into fallopian tubes and rests inside ampulla for some time. The journey time is 12-24 hours
• Human male produces 300-400 millions sperms per ejaculation. They are deposited in vagina during coitus. The process of deposition of sperms in the female genital tract is called insemination. A number of them are demobilized or eat but a number of them remain functional and undergo capacitation ( sperm activation) that provides them the ability to fertilise an ovum.
Capacitation requires 6-10 hrs.
• Capacitation consists of three process
(i) Neutralisation of inhibitory factors present in semen
(ii) Weakening of covering membrane of acrosome head by dissolution of cholesterol
(iii) Entry of Ca2+ into sperms which changes sperm movement from undulation to whiplash motion
• The activated sperm being to pass into uterus and from there to oviducts. Viscous fluid secreted by female genital tract further enhances sperm motility. A number of sperms reach the ampulla part of oviduct where the egg rests temporarily
• Fertilization involves following steps:
(i) Approximation of sperm and ovum
Sperm can remain motile for 24-48 hrs. They swim at the rate of 1.5-3 mm/min. They are able to reach the ampulla part of female genital tract partly by contraction of uterus and fallopian tubes stimulated by prostaglandins (in male semen) and oxytocin (often formed in females).
The movements are powerful within 5 minutes. After reaching an ovum, one sperm comes to lie against it. It releases lysine from its acrosomal region. Hyalouronidase and corona penetrating enzymes as well as dissolves cells of corona radiata. The sperm head now reaches zona pellucida where receptors protein fertilizing helps in attachment to specific protein of sperm. It is compatibility reaction.
(ii) Acrosome reaction
In contact with zona pellucida, acrosome covering degenerates. The contained enzymes are released. Acrosin or zona lysine dissolves zona pellucida in area of contact.
(iii) Egg reaction
A small protuberance or fertilization cone develops from the surface of ovum in the region of animal pole.
(iv) Penetration of sperm
Sperm head established contact with the lateral surface of fertilization cone. It produces a weak depolarization and ca2+ wave. Plasma membranes of the two, dissolve. Contents of head, neck and middle sperm enter ooplasm. Tail is left outside. Fertilization cone subsides. Cortical granules are extruded. They convert plasma membrane and deactivate sperm receptor of zona pellucida. A perivitelline space is created between it and zona pellucida. This prevents entry of a second sperm.
(v) Activation of ovum
Ovum undergoes meiosis II and extrudes a secondary polar body. It is now the actual ovum or female gametes.
(vi) Fusion of sperm and egg
The envelope of the sperm and egg pronuclei degenerates to form ‘synkaryon’. The act is called karyogamy or syngamy. The proximal centriole brought by sperm helps form the spindle for the division of synkaryon (cleavage nucleus). Fertilized egg is also called zygote. It immediately begins cleavage.
SIGNIFICANCE OF FERTILIZATION
(i) It restores the diploid number of chromosomes, characteristics of species i.e. 46 in human being
(ii) Fertilization initiates cleavage
CLEAVAGE
• Cleavage is a series of rapid mitotic divisions of the zygote, characterized by absence of growth of daughter cells, which convert the single celled zygote into a multicellular structure called blastula ( blastocyst)
• Cleavage differ from mitosis in the respect that
(i) There is no growth phase between successive division.
(ii) The size of cells gradually decreases
(iii) The metabolism becomes fast
(iv) There is rapid DNA replication
(v) High consumption of oxygen
• Types of cleavage
(i) Holoblastic: When whole of the egg is divided, it is found in microlecithal and mesolecithal egg. It may further be..
(a) Equal – When both the blastomers are equal Example – Amphioxus
(b) Unequal – When the blastomers are unequal in size. Example- frog
(ii) Meroblastic: When a part of the egg is divided. It is found in polylecithal eggs. It may be discoidal (e.g. birds) or superficial ( e.g. insects)
• Planes of cleavage include meridional, vertical, equatorial and transverse. Patterns can be radial ( sponges, coelenterates, some echinoderms like star fish) biradial, spiral ( flatworms, annelids, non-cephalopod mollusks) bilateral ( nematods, cephalopods, fishes, amphibians, reptiles, birds) and rotational ( placemental mammals)
MORULA
Early cleavage produces a solid ball of cells called morula
BLASTULA
• Multicellular ball like embryo produced at the end of cleavage and usually having a fluid filled blastocoels, is called blastula
• It is of the following types
(i) Stereoblastula (solid blastula): It is blastula without blastocoels. E.g. Nereis
(ii) Coeloblastula : A blastula with a prominent blastocoels. e.g. frog
(iii) Discoblastula : A blastula having a many layered disc of blastomeres above the yolk. It develops as a result of meroblastic divisions in polylecithal eggs.
E.g. Hen
(iv) Superficial blastula ( Periblastula) : A blastula having a single layer of blastomeres around the central yolk. E.g. insects
GASTRULATION
• Sum total of all the processes which convert a solid or hollow ball of cells or many layered disc of blastula into two or three germinal layers of gastrula is called gatrulation
• Gastrulation takes place by the migratory or formative or morphogentic movements of blastomers from the surface of blastula to the proper position in the gastrula.
•These movements are classified into
(a) Epiboly: Growth of one part over another like prospective ectoderm over the rest except blastocoels.
(b) Emboly: Morphogenetic movements like migration of ectoderm, mesoderm and notochord cells from surface to interior. The emboly may occur by way of
(i) Involution : Rolling of cells into interior
(ii) Invagination : Infolding
(iii) Ingression: New cells migrating into blastocoels
(iv) Delamination : Formation blastocoels is obliterated and a new cavity archenteron is formed which in future alimentary canal of the animal. Blastopore is opening of archenteron.
ORGANOGENESIS
The development of tissues and organs from the three germ layer is called organogenesis
MORPHOGENESIS
The assumption of shape, size and other morphological features by embryo is called morphogenesis
DIFFERENTIATION
It is the formation of different types of cells, which become different in size, form, chemical composition and perform different functions
FATE OF THREE GERMINAL LAYERS
Ectoderm
Central nervous system, nerves, retina, lens, cornea of eyes. Conjunctiva, ciliary and iridial muscles, lining of nasal chambers. Labyrinth, epidermis, cutaneous, glands, hair, nails, claws, hypophsis, adrenal medulla, salivary glands and enamel of teeth
Mesoderm
Dermis of skin, connective tissue, muscles, notochord, skeleton, blood, heart, blood vessel, adrenal cortex, urino – genitals system except part of urinary bladder, lining of coelom, spleen and eyes.
Endoderm
Digestive glands, liver, pancreas, middle ear, Eustachian tubes, lining of urinary bladder, respiratory system, adenophypophysis thymus, parathyroid and thyroid glands, lining of vagina and urethra, prostate.
FOETAL MEMBRANES
They are extra embryonic membranes that provide protection and nourishment to foetus. Foetal membranes are of four types
(i) Chorion – Outer foetal membrane that also takes part in formation of placenta.
(ii) Amnion – Inner foetal membrane that invests the embryo and forms a space called amniotic cavity. It is filled with fluid called amniotic fluid. Amnion protects the foetus from shock
(iii) Allantois – sac – like, develops from gut of embryo, supplies blood vessels to placenta. In reptiles and birds, it helps in respiration nutrition and excertion
(iv) Yolk sac – Membraneous sac attached to embryo near allantois, having yolk in egg laying animals and forms corpuscles in mammals till liver takes over
EMBRYO FORMATION IN HUMAN BEINGS
MORULA FORMATION
• Soon after fertilization, the zygot begins cleavage or segmentation. Cleavage consists of early mitotic divisions of fertilized egg without involving growth of daughter cells. There is rapid synthesis of new DNA and increased oxygen consumption. Surface –volume and nucleo-cytoplasmic ratio increases. The cells formed after cleavage are called blastomers. Cleavage is simple and holoblastic in humans as there is no yolk.
• The first cleavage is animal – vegetal axis or primary axis. It is slow and is completed within 30 hours of pertilization
• One of the two blastomers, is however slightly layer. Hnece, the first cleavage is holoblastic and unequal. Second cleavage is at right angle to the first one. It takes about 30 hours and completed slightly earlier in the larger blastomere so that a transitional 3-celled stage appers. Subsequent divisions are rapid and occur in different planes. They produce a solid ball of blastomers called morula. Phase of compaction ensues in 8-celed stage. Morula has almost the same size as the of fertilized egg due to presence of zona pellucida
• Morula has 16-32 cells. Th cells are compacted and of two types, outer slightly smaller peripheral cells with tight junctions than the inner mass of cells with ggap junctions. During the cleavage oung embryo descends in the fallopian tube slowly due to feeble fluid produced by epithelial secretion and cilia. Ultimately it reach the uterus. It takes 4-6 days. Corona radiate dissolved away during this period.
BLASTULATION
• In uterus, the endometrial cells become full of nutrients which are also secreted into uterine cavity. As the young embryo reaches uterus, its outer cells begins to absorb nourishment and grow while covered by zona pellucida
• The outer cells enlarge, flatten and form trophoblast. Trophoblast pours as fluid towards interior producing a cavity called blastocoels or blastocyst. It is equivalent to blastula of othe animals. The size of blastocyst is roughly three times the size of morula.
• Trophoblast then separates from inner cells except at one point called embryonic pole. The inner cells now occur at one side and called inner cell now occur at one side and called inner cell mass or embryonal knob as the latter is to form the body of embryo. Trophoblast cells in contact with inner mass are called cell of Rauber. Embryonic pole is also called animal pole. The opposite end of blastocyst is called abembryoic pole
• Blastocyst stage is completed after abour 5 days of fertilization. Trophoblast later becomes two – layered, outer synctiotrophoblast and inner cytophoblast. It secretes hCG ( human chorionic gonadotrophin). Forms villi for implantation and later on produces chorion, amnion and foetal part of placenta
IMPLANTATION
• It is a choring or embedding of blastocyst into endomerium of uterus. Implantation begins about 7th day after fertilization of ovum. It takes about process to be completed
• Blastocyst comes in contact with the endometrium in the region of embryonal knob or embryonic disc
• The surfacea cells of trophoblast secrete lytic enzymes which causes corrosion of endometrial linning. They also give rise to figure – like outgrowths called chorionic villi and uterine tissue becomes interdigitated. Villi not only help in fixation but also absorption of nourishment
• The blastocyst sink in the pit formed in endometrium then get completed buried in the endometrium that grows around it. The embedded blastocyst forms villi all around it to obtain nourishment.
• Implantation causes nutrient enrichment, enlargement of cells and vascular endometrium. Vascular endometrium ( deciduas of pregnancy) are stomal cells which have accumulated glycogen and lipid in their distended cytoplasm. The interglandular tissue increases in quantity.
• They may offer nutrition which is engulfed by the syntial trophoblast but they have been regulated as defensive mechanism
• Trophoblast covering secretes hormone called human chorionic gonadotropin ( hCG). The hormone can be detected in the urine of woman within day after implantation
• hCG maintain the corpus luteum beyond its normal life. It continues to secrete pregnancy which prevents menstruation and maintains uterine lining in nutrient rich state.
• Progesterone induces cervical glands to secrete viscous mucus for filling the cervical canal to form a protective plug. By the 16th week of pregnancy, placenta produces enough progesterone and the corpus luteum regresse.
PLACENTA
• Placenta is a temporary organ found only in mammals during gestation period and it is composed of cells derived from two different organisms, the foetus and mother
• Placenta is connection between the foetal membranes and uterine wall.
Formation of placenta
• The outer surface of the chorion in human develops a number of finger like projections known as chorionic villi, which grow into the tissue of the uterus. These villi, penetrate the tissue of the uterine wall in which they are embedded to make up the organ known as placenta by means of which the developing embryo obtains nutrients, oxygen and gets rid of carbon dioxide and metabolic wastes
• A fully formed human placenta is reddish – brown disc. It foetal surface is smooth and has the umbilical cord. The allantois gives rise to umbilical cord. Umbilical cord has two umbilical arteries (small diameter) and two umbilical veins ( large diameter)
• Umbilical arteries convey oxygen poor blood from the foetus to placenta and umbilical veins carries oxygen rich blood from the placenta to the foetus
• The blood of foetus in the capillaries of the chorionic villi comes in close contact with the mothers blood in the tissue between the villi, but are always separated by a membrane, through which substances must diffuse
• The maternal and foetal blood are not in direct contact in the placenta because
(i) Two may be incompatible
(ii) The pressure of maternal blood is far too high for the foetal blood vessels
(iii) There must be a check on passage of harmful material into foetal blood
Functions of placenta
(i) Nutritive organs – Food materials from the mother’s blood into the foetal blood through the placenta
(ii) Digestive organ – Trophoblast of placenta digests proteins before passing them into the foetal blood.
(iii) Respiratory organ – Oxygen diffuses from the maternal blood into the foetal blood through the placenta. Carbon dioxide diffuses from the foetal blood into the maternal blood also through the placenta for elimination by the mother’ lungs. Foetal haemoglobin has a greater affinity for oxygen than adult haemoglobin
(iv) Excretory organ – Nitrogenous waste such as urea, pass form the foetal blood into the maternal blood via placenta for elimination by mother’s kidney
(v) Endocrine organ – Placenta secretes some hormone such as estrogens, progesterone, human chorionic gonadotropin ( hCG), human placental lactogen ( hPL) chorionic thyrotropin, chorionic corticotrophin and relaxin.
The hCG stimulates and maintains the corpus luteum to secrete progesterone until the end of pregnancy. The hPL stimulates the growth of the mammary glands during pregnancy. Relaxin facilitates parturition by softening the connective tissue of pubic symphysis
(vi) Storage organ – The placenta stores glycogen for the foetus before liver is formed
PARTURITION
• It is the process of giving birth to a baby . The physical activities in parturition like uterine and abdominal contractions dilation of cervix and passage of baby are collectively called labour
• Labour is accompanied by localized sensation of discomfort or agony called labour pains.
• Parturation is controlled by complex neuroendocrine mechanism. Signals originate from fully formed foetus and placenta. They cause mild uterine contraction called foetal ejection reflex. It is accompanied by rise in estrogen to progesterone ratio, increase in oxytocin receptors in uterine muscles, increase in level of oxytocin secretion by both mother and foetus and stretching of uterian musculature
(i) Dilation stage
Uterine contractions begin from top. They occur once every 30 minutes. Contractions forces the baby towards cervix. The intervals between successive contraction decreases about every 1-3 minutes. Contractions are accompanied by pain caused by compression of blood vessels and uterine muscles Oxytocin induce contraction and more oxytocin secretion. The strength of uterine contraction continues to increase due to stimulatory reflex. As the baby is pushed down in uterus, its head come to lie against cervix which therefore gets dilated and stretched, A similar dilation also occurs in vagina. The first stage of labour continues for 6-12 hours. It culminates in rupturing of amniotic membrane. The amniotic fluid flows out
(ii) Expulsion stage
The intensity of uterine and abdominal contractions increases. The foetus passes out through cervix and vagina with head in forward direction in normal deliveries. It causes intense labour pain. Expulsion stage takes about 20-60 minutes. Umbilical cord is tied and cut off close to navel
(iii) After birth
Within 10-45 minutes of the delivery of baby the placenta separates from uterus and is expelled out due to series of strong uterine contractions In neonate there is a change in respiratory and circulatory system. The switch over is initiated by gaseous hormone nitric oxide (NO). Lungs expand and infant starts breathing. Blood flow through umbilical cord, formen ovale ceases. It starts passing through heart, aorta and pulmonary arteries
GESTATION PERIOD IN SOME MAMMALS
IMPORTANT DEVELOPMENTAL CHNAGES IN HUMAN EMBRYO
1. Week 1
• Fertilization
• Cleavage starts about 24 hours after fertilization
• Cleavage to form a blastocyst after fertilization
• More than 100 cells
• Implantation 6-9 days after fertilization
2. Week 2
• The three primary germ layers develops
3. Week 3
• Woman will not have period. These may be first sign of pregnancy
• Beginnings of backbone
• Neural tube develops, the beginning of brain and spinal cord
4. Week 4
• Heart, blood vessels, blood and gut start forming
• Umbilical cord developing
5. Week 5
• Brain developing
• ‘Limb buds’ a small swelling which are the beginnings of arms and legs
• Heart is a large tube and starts to beat, pumping blood. This be seen on a ultra sound scan
6. Week 6
• Eyes and ears start to form
7. Week 7
• All major internal organ developing
• Face, forming
• Eye have some colour
• Mouth and tongue develop
• Beginnings of hands and feet
• 2.5 cm long
8. By week 12
• Foetus fully formed with all organs, muscles, bone, toes and fingers
• Sex organs well developed
• Foetus reaches 7.5cm in height and about 14g weight
9. By week 20
• Hair beginning to grow, including eyebrows and eyelashes
• Fingerprints developed
• Fingernails and toenails growing
• Firm hand grip
• Between 16 and 20 weeks bay usually felt moving for first time
10. Week 24
• Eyelids open
• Foetus measures about 32 cm and weighs about 650gm
11. Week 26
Has a good chance of survival if born prematurely
12. By week 28
• Baby moving vigorously
• Responds to touch and noise
• Swallowing amniotic fluid
13. By week 30
Usually lying head down ready by birth
Foetus is about 43cm long and its weight is about 1800gm
14. By week 40
Birth
Generally child is about 50cm long and weighs about 3300gm
LACTATION
• Production of milk in the female’s breasts following the birth of a young one in mammals is called lactation
Preparation of breast ( mammogenesis)
During pregnancy, the breast enlarges due to growth of mammary glands Synthesis and secretion from the breast alveoli ( lactogenesis) Secretion and storage of milk begins after birth of the young one, usually within 24 hours under the influence of prolactin
When the estrogen and progesterone are withdrawn following delivery, prolactin begins its milk secretory activity in previously fully developed mammary glands
Ejection of milk
The actual release of milk called milk letdown, requires the presence of oxytocin, which brings about contraction of smooth muscles of the ducts within the mammary gland. Secretion of prolactin and oxytocin depends on suckling stimuli produced by the nursing infant on the nipples of breasts Maintenance of lactation ( galactopoiesis)
For maintenance of lactation, suckling is important
Milk pressure reduces the rate of production and hence periodic breast feeding is necessary to relieve the pressure which is in turn maintains the secretion.
• After birth, the breast first release is not milk, but colostrums for 2 or 3 days. It is a thin, yellowish, fluid called foremilk which is rich in protein, antibodies but low in fat.
• Human milk consists of water, fat, casein, lactose, mineral salts and vitamins. A nursing woman secretes 1 to 2 litres of milk per day
DEVELOPMENTAL DISPRDERS
AMNIONITIS
It is inflammation of amnion, usually resulting from premature rupture of amnion
ABORTION
It is giving birth to an embryo or foetus at the stage of about 20 weeks of gestation
TERATOGENY
Teratogens are certain agents or drugs that cause abnormal development in developing embryo/foetus. It may cause malformation in developing embryo.
ECTOPIC PREGNANCY
• The developmental site of foetus is other than the uterus like fallopian tube or cervix
• The growth of foetus may cause tube to rupture and bleed and my lead to miscarriage
• The condition is diagnosed by ultrasound and foetus may be removed by laproscopy before damage is done to fallopian tube.