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1.ESC/00176/022 ELVIS OKETCH
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Cleavage in birds is a crucial early developmental process that follows fertilization and leads to the formation of a multicellular embryo. It occurs under specific structural and functional adaptations to support the development within the large, yolk-rich egg. Here’s a detailed discussion of the cleavage process in birds, its unique characteristics, and how it contributes to embryogenesis:
1. Overview of Bird Egg Structure
Bird eggs are telolecithal, meaning they contain a large quantity of yolk concentrated at one end, the vegetal pole, while the animal pole contains the blastodisc, a small, yolk-free region where fertilization and cleavage occur. The yolk serves as the nutrient source for the developing embryo, and its abundance poses unique challenges for cellular division, as the entire egg cannot divide due to the yolk’s volume and density.
2. Meroblastic (Partial) Cleavage in Birds
Birds exhibit meroblastic cleavage, which means that only part of the egg undergoes division. In birds, this is specifically a discoidal meroblastic cleavage, where cleavage is confined to the blastodisc at the animal pole. The yolk itself does not divide, and cellular divisions are restricted to a small, disc-shaped region atop the yolk.
3. Cleavage Process
The cleavage process in bird embryos is typically divided into several stages as follows:
a. First Cleavage Divisions
Initial Divisions: Cleavage begins within the blastodisc, with the first division being vertical, producing two daughter cells within this small region. The second cleavage is also vertical and at a right angle to the first, resulting in four cells. These initial divisions create a limited number of cells, which remain attached to the underlying yolk.
Subsequent Divisions: The third cleavage is horizontal (parallel to the surface), creating an eight-cell stage. Further divisions continue primarily within the blastodisc, forming a multilayered cluster of cells known as the blastoderm.
b. Formation of the Blastoderm
As cleavage progresses, the blastoderm expands across the blastodisc. The cells in the blastoderm undergo incomplete cytokinesis, so they remain loosely connected at the yolk surface.
With continued cleavage, a separation begins between the blastoderm and the yolk beneath it. This creates a fluid-filled space called the subgerminal cavity, which helps define the blastoderm as a distinct structure.
Eventually, the blastoderm differentiates into two main layers: the epiblast, which will give rise to the embryo, and the hypoblast, which contributes to extraembryonic structures.
4. Epiblast and Hypoblast Formation
The blastoderm reorganizes into two essential layers:
Epiblast: The upper layer that will form all three primary germ layers (ectoderm, mesoderm, and endoderm) through the process of gastrulation. This layer undergoes further specialization to develop into the embryo proper.
Hypoblast: The lower layer derived from some cells of the blastoderm and others migrating from the edge of the blastoderm. The hypoblast does not contribute to the embryo itself but instead forms structures essential for early development, such as the yolk sac.
Together, the epiblast and hypoblast layers create the bilaminar blastoderm, which will guide further embryonic development.
5. Development of the Area Pellucida and Area Opaca
Two distinct regions emerge within the blastoderm:
Area Pellucida: The central, clear region of the blastoderm directly above the subgerminal cavity. This region is where most embryonic cells will form.
Area Opaca: The peripheral area surrounding the area pellucida, where the cells remain in contact with the yolk and help provide nutrients to the developing embryo.
6. Formation of the Primitive Streak and Gastrulation
Once cleavage completes, the next stage is gastrulation, a key developmental event where cells start moving and differentiating to form the basic body plan:
Primitive Streak: The formation of a groove called the primitive streak along the midline of the epiblast signals the start of gastrulation. Cells from the epiblast move through this streak, migrating to specific regions to form the three primary germ layers.
Germ Layer Formation: Through the primitive streak, epiblast cells dive inward, creating the mesoderm and endoderm, while cells remaining on the surface form the ectoderm. These germ layers will later give rise to all tissues and organs of the bird.
7. Special Characteristics and Adaptations in Bird Cleavage
Cleavage in birds is adapted to their yolk-rich eggs, differing significantly from mammals and amphibians:
Discoidal Cleavage: Restricted to a small area due to the large yolk volume, which would otherwise inhibit complete cleavage.
Meroblastic Nature: Allows the yolk to remain intact, providing a continuous nutrient supply while the blastoderm develops.
Nutritional Strategy: The large yolk mass supports the bird embryo’s entire development inside the egg, as opposed to placental mammals where the embryo receives nutrients from the mother.
8. Significance of Cleavage in Bird Development
The cleavage stage sets up the foundational structure for the embryo and establishes cell layers and spatial organization that are critical for subsequent stages of development:
Establishing Body Axes: The primitive streak formed during gastrulation also helps in defining the embryo’s anterior-posterior axis.
Supporting Self-Sustained Development: By forming layers and cavities like the subgerminal cavity, cleavage in birds prepares the embryo for nutrient absorption and protection within the egg.
Conclusion
Cleavage in birds is a unique process adapted to the telolecithal structure of their eggs, enabling the embryo to develop within a nutrient-rich yolk environment. This process of discoidal meroblastic cleavage leads to the formation of the blastoderm, which further differentiates into structures essential for forming the body plan of the bird. The complexity of cleavage in birds provides insights into how vertebrates adapt their reproductive strategies to their environments and developmental needs, ensuring the successful growth and emergence of the bird embryo.
References: Here are some key references for studying cleavage in birds:
1. Gilbert, S. F., Barresi, M. J. F. (2016). Developmental Biology (11th ed.). Sunderland, MA: Sinauer Associates.
This textbook provides an in-depth exploration of developmental biology principles, including cleavage and gastrulation processes in various organisms, such as birds.
2. Patten, B. M. (1971). Foundations of Embryology (3rd ed.). New York: McGraw-Hill.
Patten’s work is a classic reference that covers the foundational concepts of embryology, including cleavage patterns in birds and other vertebrates.
3. Romanoff, A. L. (1960). The Avian Embryo: Structural and Functional Development. New York: Macmillan.