Development

Chapter 4

Definition

"the gradial changes taking place in a fertilized egg leading to formation of a living being, which can survive independantly and can change into an adult are called Development."

Embryology

"The branch of biology in which changes leading to formation of an independent individual ffrom a fertilized egg are studied is called Embryology."

Egg

Egg is an apparatus containing female gamete. For nourishment of developing embryo, it contains a nutritional compound called yolk. Yolk is a complex mixture of proteins, lipids (fats), carbohydrates, vitamins and minerals.

Types of Eggs

Eggs are classified according to the amount of yolk present in them. Different categories of eggs are as follows:

  1. Isolecithal Egg

    These are the eggs having small amount of yolk mixed homogenously with the cytoplasm. These are of two types:

    (i) Alicethal Egg

    These are the eggs, which are devoid of yolk or may contain a negligible amount of yolk, for example eggs of Eutherians.

    (ii) Meiolicethal Eggs

    These are the eggs containing small or small amount of yolk. For example eggs of Amphibians.

  2. Telolicethal Egg

The egg containing moderate to large amount of yolk. These eggs are of two types:

(i) Mesolicethal Egg

These eggs contain moderate to large amount of yolk which generally concentrates towards the lower half or vegetal pole of the egg. For example eggs of Amphibians.

(ii) Polylicethal Eggs

These eggs have got a large or enormous amount of yolk generally separated from egg’s white. For example eggs of birds and reptiles.

= Structure of Frog =

A single female frog lays hundreds of eggs in water which are held together by a jelly like substance. The cluster thus form is called Frog’s Spawn.

A single egg of frog is round or spherical measuring about 1to2 millimeters in diameter. It is externally covered with three gelatinous layers. These transparent layers act as an effective convex lens to converge maximum sunlight to produce necessary temperature beneath these layers lies the vitelline membrane along with Perivitelline Space. The outer and superior or upper half portion of egg is pigmented (containing melanim) appears black, while the lower half is whitish in colour. The black pigment serves to absorb sunlight to produce necessary heat during Embryo Genesis. The black half of egg contains more cytoplasm than the lower yolk containing half. For this reason upper half is metabolically active and called Animal Pole. The lower yolk containing half is called Vegetal Poles. The egg as a whole is a single cell.

= Fertilization in Frog =

Fertilization

"Fusion of male and female pronuclei or simply fusion of male and female gametes is called fertilization."

Process in Frog

Female frog lays egs in water over which the male frog sheds sperms. As a rule a single sperm is capable of fertilizing a single egg. When a sperm touch the egg, it causes some physiological change in the egg by which the egg does not permit any other sperm to enter. Now, the head of the sperm bores its way and enters into the egg leaving its tail behind and outside. Here the sperm head becomes rounded and now called male pronucleus after completing into second meiotic division. Similarly the nucleus of egg completes its 2nd meiotic division and forms female pronuclei. Now the two haploid pronuclei unite to form zygote nucleus which is Diploid (2n) in nature. Thus fertilization is completed.

After fertilization the pigment of the egg moves upward on the side lying opposite to the Reception Cone i.e. the point of entry of sperm in the egg, the bare area thus formed is called The Grey Crscent extending up to half way around egg.

Division of Zygote

  1. Cleavage

    The zygote begins to divide 2 to 3 hours after fertilization. The initial cell division is called Cleavage.

    The first cleavage is vertical, starting from the animal pole extending up to vegetal pole. Thus zygote divides into two equal and similar cells called Blastomeres. This type of complete division is called Holoblastic Division.

    The second division takes place soon afterwards (30 to 45 min later). It is also vertical but at right angle to the first division forming 4 equal and similar Blastomeres.

    The third cleavage is horizontal and above the equator, dividing 4 blastomeres into 8 cells out of which upper 4 cells are smaller and contain more cytoplasm. The lower 4 cells are towards the vegetal pole, they are larger and contain more yolk. These cells are called Micromeres or Megameres respectively. Later on two simultaneous division take place vertically by which 16 cells are formed . Two horizontal divisions (one nearer to each pole) now divide the cells into 32. Further cleavage becomes irregular as cytoplasmic micromeres divide more rapidly.

  2. Morula

    As a result of rapid cell division, mulberry shaped ball of cells is formed called Morula. It is initially solid but soon a cavity appears nearer to the animal pole known as Blastocoel.

  3. Blastula

As cell division advances, a cavity appears inside the cellular mass nearer to animal pole called Blastocoel and the hollow sphere of cells is called Blastula. Micromeres divide more rapidly than the megameres and soon overlap part of the megameres forming a band called Germ Ring. The cells lying at Gray crescent divide more rapidly than rest of the cells.

= Gastrulation =

Definition

"Movement, positioning and arrangement of cells is called Gastrulation."

Classification

The process of Gastrulation involves three types of Movements of cells namely:

  1. Epiboli
  2. Emboli
  3. Involution
  1. Epiboli

    It is the down growth of micromeres or animal cells over the yolk containing megameres. It is due to rapid proliferation of micromeres that they over grow and more down ward on all sides except a small area called Blastopore. Through this small area, megameres seem to hang outward constituting the Yolk Plug.

    Blastopore

    As a result of epiboli, a small area of megameres remains uncovered by the micromeres known as Blastopore. Initially it appears as a crescent shape area, later on it becomes horse and shoe shaped with horizontal upper margin called Dorsal Lip. It soon becomes ring shaped with a Dorsal as well as Ventral lip and protruded megameres in between the constituting yolk plug.

  2. Emboli or Invagination

    After formation of Blastopore, the animal cells linning the Dorsal lip begin to invaginate i.e. they migrate inward along the dorsal lip forming a small slit like cavity between the dorsal lip and the invaginating cells. This is the initiation of Archenteron. This cavity further deepens with time forms a definite hollow cavity.

  3. Involution

With the beginning of Emboli, the cells of the srufac eare displaced along the Dorsal lip to take their definite position for the formation of organs. This massive movement of different cells to their definite positions is called Involution.

The cells of Dorsal lip migrate and turn inward along the ceiling of archenteron forming Chorda Mesoderm.

Archenterone expands dorsally and laterally, the floor of it is made up of small yolk cells while the lateral walls are composed of micromeres. During the course of its expansion the blastocoel gradually diminishes. Megameres tend to lie along the ventral lip and floor archenteron.

Rotation

Due to shifting and distribution of cells and expansion of Archenterone with diminution in Blastocoel, the centre of gravity of Gastrula also shifts. The animal pole which was formally located on the top now comes to lie antero-ventally shifting nearly 120° .

The Germinal Layers

After completion of Gastrulation, Blastopore closes completely forming a small longitudinal groove open at the ends. The ventral opening closes and forms the anal pit while the dorsal aperture remains open and forms Nementeric Canal. At this moment the gastrula becomes Pear shaped and composed of an outer layer of micromeres called Ectoderm. The invaginating cells lining the archenterone separate out from other layers of cells to form Endoderm.

The cells between these two layers delaminate and differentiate to form Mesoderm. Along the Mid Dorsal Line the cells of Mesoderm form a slid rod like structure called Notochord. The Mesodermal cells grow on both the sides of Notochord and finally, after forming two complete sheets, meet at the Mid-Ventral Line. Thus the Embryo becomes three layered which are called Primary Germinal Layers.

Fate of Primary Layers

(i) Ectoderm

It gives rise to Epidermis, cutaneous glands, nervous system, parts of sensory organs, enamel of teeth, Internal lining of Cloaca and Mouth, Nasal Mucosa.

(ii) Endoderm

It forms mucosa of midgut, liver, spleen, pancreas, lung, urinary bladder and thyroid glands.

(iii) Mesoderm

It gives rise to body muscles, connective tissue, dermis, heart, kidneys, Reproductive organs, blood vascular system, Notochord and bones.

= Organogenesis =

Formation of bodily organs and systems by the germinal layers is called organogenesis. Few are given below:

  1. Central Nervous System

In ectodermal cells of Mid Dorsal Line becomes differentiated and form a thickened plate known as Neural Plate. The edges become much thicker and fold outward forming New Folds. These folds grow mid dorsally and converge to fuse, enclosing a canal inside known as Neural Canal. After fusion of folds, the canal is changed into a neural tube. Anteriorly the tube forms three hollow swellings, which form fore brain, mid brain and hind brain. Rest of the tube forms spinal cord. This Notocord lying below the neural tube between the folds of mesoderm gradually forms Vertebral Column.

2. Digestive System

In the elongated Embryo, invagination of ectoderm occurs below the anterior end forming a pit known as Stomodaeum, the future buccal cavity. Similarly invagination of ectoderm occurs at closed blastopore posteriorly to form Proctodaeum or the future Cloaca. Stomodaeum and Proctodium open interiorly into Archenterone forming a continuous tube. The middle portion of this tube i.e. Mesenterone differentiate into different parts forming Alimentary Canal. Buds grow outward from mesenterone at different levels, which give rise to lung, liver and pancreas. The anterior part of Mesenterone just near its junction with stomodaeum forms 5 pairs of Pharyngeal Pouches.

Pharyngeal Pouches

Pharyngeal Pouches are important embryonic structures. First pair of pouches forms the tympanic membrane, Estachian tube and middle ear. The remaining 4 pairs of pouches open to the exterior and forms external gills later on.

= Tad Pole Larva =

Within 48 hours after fertilization, the embryo becomes much developed that it can survive independently. During this period following changes take place at and after hatching.

Metamorphosis

Definition

"The phenomenon in which a living being passes through different and successive morphological and physiological changes to reach the adult stage is called Metamorphosis."

Classification

In metamorphosis of Frog changes taking place can be grouped into two types, which are given below:

  1. Morphological Changes
  2. Physiological Changes

1. Morphological Changes

"The changes taking place in the shape and appearance of Tadpole are called morphological changes. "

The two main types of Morphological Changes are:

  1. Regressive Changes
  2. Progressive Changes
  1. Regressive Changes
  1. Progressive Changes

2. Physiological Changes

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