Chapter 5: Life History & Early Embryology

Life History and Early Embryology

Introduction to Life History

● Definition
○ Life history refers to the sequence of events in an organism's life from fertilization
to death.
● Philosophical Perspective
○ Quote by Benjamin Disraeli: “Youth is a blunder; manhood a struggle; old age a
regret.”
■ Implication for biologists: Life history encompasses development stages
rather than emotional or moral interpretations.

Stages of Life History

● A. Fertilization
○ Process
■ Union of sperm (male gamete) and ovum (female gamete).
■ Results in a diploid zygote.
● B. Embryonic Development (Ontogeny)
○ Overview
■ Extends from fertilization to birth or hatching.
○ Key Stages:
■ Cleavage: Rapid cell divisions leading to a multicellular structure.
■ Blastula: Formation of a hollow sphere of cells with a fluid-filled cavity
(blastocoel).
■ Gastrulation: Rearrangement of cells into three germ layers.
■ Organogenesis: Development of organs from germ layers.
● C. Maturation
○ Definition
■ The phase from birth to sexual maturity, involving:
■ Growth in size.
■ Development of reproductive anatomy and learned behaviors.
○ Terminology
■ Juveniles or immatures: Pre-reproductive individuals.
 ● D. Senescence
○ Definition
■ The process of aging, characterized by:
■ Loss of physical vigor.
■ Decline in reproductive capability.
○ Examples
■ Rare in wild animals; more common in captive settings (e.g., zoos).
■ Notable wild examples: salmon that senesce after spawning.

Senescence and Aging

● A. Overview of Senescence
○ General patterns of aging and vulnerability to predation in the wild.
○ Human society’s unique approach to aging individuals.
● B. Human Aging
○ Differences from other species:
■ Extended post-reproductive life.
■ Possible roles of elderly: caregivers, repositories of knowledge.

Early Embryology

● A. Germ Layer Formation

○ Three Primary Germ Layers:
■ Ectoderm: Develops into skin and nervous system.
■ Mesoderm: Forms muscles, bones, and circulatory system.
■ Endoderm: Develops into digestive and respiratory systems.
○ Homology: Similar embryonic development across species suggests evolutionary
relationships.
● B. Stages of Embryonic Development
○ Zygote: Fertilized egg.
○ Morula: Solid ball of cells after several rounds of cleavage.
○ Blastula: Hollow sphere of cells; important for subsequent gastrulation.
○ Gastrula: Formation of germ layers and initial body plan.
○ Neurula: Development of the nervous system, including neural tube formation.

Fertilization Process

● A. Definition of Fertilization
○ Activation of embryonic development upon sperm penetration.
● B. Ovum Structure
○ Types of Yolk:
■ Microlecithal: Minimal yolk (e.g., some fish).
 ■ Mesolecithal: Moderate yolk (e.g., amphibians).
■ Macrolecithal: Abundant yolk (e.g., birds).
○ Polarity:
■ Vegetal pole (yolk-rich area).
■ Animal pole (nucleus-rich area).
● Ovum Envelopes
○ Primary Egg Envelope: Includes vitelline membrane (analogous to zona pellucida
in mammals).
○ Secondary Egg Envelope: Follicle cells that assist in nutrient transfer.
○ Tertiary Egg Envelope: Forms during passage through the oviduct (e.g., shells in
birds).

Reproductive Strategies

● Oviparity vs. Viviparity

○ Oviparous: Laying eggs (e.g., birds, reptiles).
○ Viviparous: Live-bearing; embryo develops inside the mother.
● Distinction Between Parturition and Oviposition
○ Parturition: Act of giving birth.
○ Oviposition: Act of laying eggs.
● Fetal Nutrition Types
○ Lecithotrophic: Nutrient supply from yolk.
■ Examples: Some fish embryos.
○ Matrotrophic: Nutrients supplied by the mother, either through a placenta or other
means.
■ Examples: Mammals.

Nutritional Strategies for Embryos

● Lecithotrophic Nutrition
○ Direct transfer of nutrients from yolk to embryo.
● Matrotrophic Nutrition
○ Nutrients provided through:
■ Vascular placentae.
■ Secretions from the oviduct.
○ Postparitive Matrotrophy: Continued nutrient delivery through lactation in
mammals.
Cleavage

Cleavage is a critical phase in early embryonic development characterized by repeated mitotic

cell divisions of the zygote. During this stage, the embryo undergoes minimal growth in size,
transitioning from a single cell into a solid mass known as the morula, and eventually forming a
multicellular and hollow structure called the blastula.

1. Formation of Blastomeres

● The initial cleavage events produce cells known as blastomeres, resulting from the
divisions of the ovum. The first cleavage furrows emerge at the animal pole and extend
toward the vegetal pole.
● In species with minimal yolk, such as amphioxus and eutherian mammals, cleavage is
holoblastic, meaning the mitotic furrows traverse the entire zygote. This results in a
complete division of the egg.
● As cleavage continues, the structure forms a hollow ball of cells surrounding an internal
fluid-filled cavity called the blastocoel.

2. Types of Cleavage

● Holoblastic Cleavage: Occurs in eggs with little yolk, leading to complete division of
the zygote. In amphioxus, for example, this leads to the formation of a 32-celled blastula.
● Meroblastic Cleavage: Characteristic of yolk-rich eggs, where cell division is limited. In
cases like many fish, reptiles, and birds, only a portion of the egg cytoplasm divides,
often resulting in discoidal cleavage where cleavage occurs in a disc shape at the animal
pole, leaving the yolk undivided at the vegetal pole.

3. Species-Specific Examples of Cleavage Patterns

● Amphioxus: Exhibits microlecithal eggs with holoblastic cleavage. The initial cleavage
planes lead to a four-celled embryo, which subsequently reaches the 32-celled blastula
stage.
● Fishes:
○ In gars and bowfins, cleavage is holoblastic but slower at the vegetal pole. Most
cell divisions occur at the animal pole, producing a blastula similar to that of
amphibians.
○ In hagfishes, chondrichthyans, and most teleosts, cleavage is discoidal, with the
majority of the yolk remaining undivided, resulting in distinct cell populations:
the blastoderm and periblast.
● Amphibians: Similar to fish, where blastomeres at the animal pole divide more
frequently, resulting in a smaller blastoderm over a large yolk-rich vegetal pole.
 ● Reptiles and Birds: Cleavage is discoidal due to substantial yolk reserves. The
blastoderm forms at the animal pole, with a subgerminal space developing between the
blastoderm and yolk.
● Mammals:
○ Monotremes: Produce macrolecithal eggs with discoidal cleavage, forming a
blastoderm that envelops the yolk.
○ Marsupials: Have modest yolk accumulation and do not form a morula. Instead,
they develop a unilaminar blastocyst, with cells differentiating into an embryo and
trophoblast.
○ Eutherians: Characterized by minimal yolk. After fertilization, a compact morula
forms, which then develops into a blastocyst with distinct trophoblastic layers,
essential for establishing maternal-fetal connections.
○ The terms choriovitelline membrane and chorioallantoic membrane are
suggested alternatives to the term trophoblast.

This distinction is significant:

● If the trophoblast is unique to eutherians:

○ It implies this structure arose as a new embryonic feature during the Cretaceous
period.
○ This emergence coincides with the rise of eutherian mammals.
○ The newly evolved trophoblast would be crucial in the eutherian reproductive
strategy, facilitating prolonged intrauterine exchange between fetal and maternal
tissues.

The trophoblast in marsupials performs many of the same functions as in eutherians.

Until more conclusive evidence emerges:

Overview of Cleavage

● Definition: Cleavage involves repeated cell divisions that produce a multicellular

structure known as the blastula.
● Cell Composition:
○ Each cell, or blastomere, contains some of the original cytoplasm of the egg.
○ Due to uneven distribution of ingredients within the polarized ovum, each cell has
a slightly different cytoplasmic composition.
● Blastula Size:
○ In some species, the blastula can absorb uterine fluids, causing it to swell in size.
○ The blastula does not grow by incorporating new cells.
● Transition to Gastrulation:
○ Following blastula formation, the next stage is gastrulation.
 ○ Most cells migrate to their final positions within the embryo during this stage.
● Cell Fate:
○ Initially, cells have a broad ability to differentiate into various cell types.
○ As development progresses, the potential for differentiation narrows.
○ By the end of gastrulation, most cells are committed to contribute to specific parts
of the embryo.
● Final Differentiation:
○ In subsequent embryonic stages, cell fate becomes increasingly restricted until
each cell differentiates into a terminal cell type.

Gastrulation

Gastrulation involves major cell rearrangements to form the three primary germ layers:

● Ectoderm (outer layer)

● Mesoderm (middle layer)
● Endoderm (inner layer)

During this stage, cells move inward from the surface of the blastula, reorganizing to establish
the gut structure (archenteron or gastrocoel) and the mesodermal body cavity (coelom). Key
methods of cell movement during gastrulation include:

● Epiboly: Spreading of cells over the surface.

● Involution: Cells turning inward and spreading.
● Invagination: Folding inward of a cell layer.
● Delamination: Splitting of a cell sheet into layers.
● Ingression: Individual cells migrating to the interior.

Neurulation

Neurulation follows gastrulation and involves the formation of the neural tube from the
ectoderm, a precursor to the central nervous system. This process occurs primarily through:

● Primary neurulation: The ectoderm thickens into a neural plate, which folds to form
neural folds that fuse to create the neural tube.
● Secondary neurulation: Observed in some species like lampreys and teleosts, where the
neural tube forms from cavitation within a solid rod of ectodermal cells (neural keel).

Germ Layer Differentiation

 ● Endoderm: Develops into the gut and associated organs.
● Mesoderm: Forms structures like the notochord, somites, and coelom, differentiating
into:
○ Paraxial mesoderm (epimere): Forms somites, contributing to muscles and
vertebrae.
○ Intermediate mesoderm (mesomere): Forms parts of the kidney.
○ Lateral plate mesoderm (hypomere): Creates the coelom, divided into
splanchnic and somatic mesoderm.

Coelom Formation

Coelom formation can occur through two mechanisms:

● Enterocoely: Primitive method where the coelom forms as the mesoderm pinches off
from other layers.
● Schizocoely: The mesoderm forms as a solid sheet and later splits.

Most vertebrates predominantly exhibit schizocoely, despite their classification as

deuterostomes, which typically feature enterocoely. This divergence has led to ongoing debates
about the evolutionary significance of coelom formation.

Neural Crest Cells

As neurulation progresses, some ectodermal cells form neural crest cells, which migrate to
various locations and differentiate into diverse structures, making them unique to vertebrates.

Gastrulation in Amphioxus

● Process: Invagination of the vegetal wall.

● Transformation:
○ Embryo changes from a single layer of blastomeres to a double layer (ectoderm
and endomesoderm).
● Endomesoderm:
○ Forms the primitive gut and the gastrocoel.
○ Communicates with the exterior through the blastopore.
● Neurulation:
○ Mesoderm delineation occurs with paired outpocketings forming the coelom.
○ Chordamesoderm:
■ Differentiates into the notochord.
■ Stimulates ectoderm differentiation into the central nervous system.
○ Lateral epidermis moves to cover the neural plate, allowing the neural tube to
form beneath.
Gastrulation in Fishes

● Variation: Patterns depend on yolk content.

● Lampreys and Primitive Bony Fishes:
○ Begin with a dorsal indentation at the blastopore.
○ Surface cells move inward via epiboly to form the endomesoderm around the
gastrocoel.
● Sharks and Teleost Fishes:
○ Blastoderm expands over the yolk, forming the extraembryonic yolk sac.
○ Endomesoderm arises beneath and at the edges of the blastoderm.
○ Thickest at the posterior edge, forming the embryonic shield.
● Endoderm and Mesoderm:
○ Endoderm becomes a flat sheet over the yolk.
○ Mesoderm forms chordamesoderm at the midline, contributing to the notochord
and lateral mesodermal plates.
○ Yolk enclosed by periblast, mesoderm, and ectoderm (no endoderm).

Coelom Formation

● Lampreys: Coelom is enterocoelic (primitive method).

● Most Vertebrates: Coelom forms by schizocoely (mesoderm splits to create cavities).

Gastrulation in Amphibians

● Initiation: A superficial indentation marks the beginning, establishing the dorsal lip of
the blastopore.
● Major Cell Movements:
○ Epiboly: Surface cells move towards the blastopore from all directions.
○ Involution: Cells involute over the lips of the blastopore.
○ Migration: Entering cells settle into specific positions within the embryo,
forming the endomesoderm surrounding the gastrocoel.
● Chordamesoderm Formation:
○ Arises middorsally within the endomesoderm and will become the notochord.
● Separation of Germ Layers:
○ Paired projections grow out from the lateral inner wall of the endomesoderm and
meet beneath the forming notochord.
○ These projections and the ventral region of yolk-laden endomesoderm separate to
form:
■ Endoderm proper
■ Mesoderm proper (rest of the endomesoderm)
● Mesoderm Development:
 ○ Grows downward between the delineated endoderm and outer ectoderm.
○ Distinct regions emerge:
■ Epimere (paraxial mesoderm)
■ Mesomere (intermediate mesoderm)
■ Hypomere (lateral plate mesoderm)
● Coelom Formation:
○ Occurs by schizocoely, where the solid mesodermal layer splits to create the
coelom.
● Key Outcomes of Gastrulation:
○ Establishes ectoderm and endoderm.
○ Obliterates the blastocoel and forms the new gastrocoel.
○ Leaves a partially plugged blastopore with yolk-laden cells.

Neurulation in Amphibians

● Timing: Begins before the endomesoderm has fully separated into distinct germ layers.
● Process:
○ Involves thickening of the neural plate, which rolls up to form the hollow neural
tube.

Gastrulation in Birds

● Embryonic Structure:
○ Flattened blastula: Composed of:
■ Epiblast (superficial layer)
■ Hypoblast (beneath the epiblast)
■ Blastocoel (space between them)
● Initiation:
○ Marked by the appearance of a thickened area in the epiblast, forming the
primitive streak.
○ Primitive node (Hensen’s node) originates at the posterior region of the embryo.
○ Primitive groove: A narrow gully along the primitive streak.
● Cell Movements:
○ Epiboly: Cells spread across the surface of the epiblast towards the primitive
streak.
○ Involution: Cells involute at the edges of the primitive streak and enter the
embryo.
■ Mesoderm: Formed by cells spreading between the epiblast and
hypoblast.
■ Endoderm: Formed by cells sinking to the level of the hypoblast,
displacing hypoblastic cells outward.
 ● End of Gastrulation:
○ Surface cells from the epiblast migrate to new positions.
○ Remaining surface cells form the ectoderm.
○ Involuting cells replace hypoblast cells with embryonic endoderm over the yolk.
○ Mesoderm forms between ectoderm and endoderm, with a notochord
differentiating along the midline.

Neurulation in Birds

● Process:
○ Formation of a neural tube from a neural plate precursor.
○ Three germ layers (ectoderm, mesoderm, endoderm) are already delineated.
○ Lateral mesoderm reorganizes:
■ Mesoderm is a solid plate with recognizable epimere, mesomere, and
hypomere.
■ Hypomere splits to form:
■ Splanchnic mesoderm (inner layer)
■ Somatic mesoderm (outer layer)
■ Schizocoelic coelom between these layers.
● Mesoderm Associations:
○ Mesodermal layers associate with adjacent endoderm and ectoderm to produce:
■ Splanchnopleure (endoderm + splanchnic mesoderm)
■ Somatopleure (ectoderm + somatic mesoderm).

Gastrulation in Reptiles

● Development: Less studied than in birds but assumed similar.

● Gastrulation:
○ Retains a discrete blastopore associated with cell movement during gastrulation.
○ Unlike birds, where the blastopore is lost, reptiles exhibit cell streaming through
the blastopore.

Gastrulation in Mammals

Monotremes

● Gastrulation:
○ Similar to reptiles; involves a disc atop a large yolk mass.
○ Blastocyst: Unilaminar at the end of cleavage; blastoderm is 5-7 cells thick at the
center, thinner at the margins.
 ○ Pregastrulation: Becomes bilaminar as the blastoderm grows around the yolk,
forming distinct endoderm (inward-moving cells) and ectoderm (surface cells).
○ Primitive Streak: Marks the beginning of gastrulation; serves as a major
organizing axis.
○ Cell Movements: Involves epiboly and involution to bring surface cells into the
embryo.
○ Mesoderm Formation: Arises from cells entering via the primitive streak,
positioned between ectoderm and endoderm.
● Neurulation:
○ Involves rolling up of the neural plate into a hollow neural tube.

Marsupials

● Gastrulation:
○ Blastocyst: Unilaminar, consisting of protodermal cells.
○ Pregastrulation: Transforms into a bilaminar embryo (ectoderm and endoderm)
as protoderm cells proliferate and migrate.
○ Primitive Streak: Appears in the ectoderm, marking the beginning of
gastrulation.
○ Cell Movements: Surface cells stream to the primitive streak and involute into
the embryo, contributing to the mesoderm between ectoderm and endoderm.
● Neurulation:
○ Neural plate rolls into a neural tube, similar to other vertebrates.

Eutherians

● Gastrulation:
○ Blastocyst: Composed of an outer trophoblast and an inner cell mass.
○ Pregastrulation: Inner cell mass reorganizes into a bilaminar embryonic disc
(epiblast and hypoblast).
■ Hypoblast forms a thin layer, sometimes referred to as endoderm.
○ Primitive Streak: Marks the start of gastrulation; epiblast surface cells stream
toward it.
○ Cell Movements: Involves epiboly and involution; some cells displace the
hypoblast, forming a middle mesoderm.
○ Notochord Formation: Arises from entering mesodermal cells.
● Neurulation:
○ Develops a tubular neural tube from the neural plate.
○ Mesoderm differentiates into epimere, mesomere, and hypomere.
○ Coelom formation occurs via schizocoely, defining somatic and splanchnic
mesodermal sheets.
Organogenesis

● Post-Neurulation Changes:
○ Transition from animal-vegetal polarity to bilateral symmetry based on the
anterior-posterior axis.
○ Three Germ Layers:
■ Ectoderm: Develops into nervous tissue and epidermis.
■ Endoderm: Forms lining of digestive and respiratory systems.
■ Mesoderm: Gives rise to skeletal, muscular, circulatory systems, and
connective tissues.
○ Germ layers are strategically positioned for interaction during organogenesis,
where two layers often combine to form organs (e.g., alimentary canal from
endoderm and mesoderm).

Histogenesis

● Definition: Tissue formation from cellular germ layers (ectoderm, mesoderm,

endoderm).
● Primary Tissue Categories:
○ Epithelial Tissue
○ Connective Tissue
○ Muscle Tissue
○ Nervous Tissue (discussed in detail in other chapters).

Epithelium Tissues

● Characteristics:
○ Composed of closely packed cells with minimal extracellular matrix.
○ Has a basal layer (formerly called basement membrane) and a free surface
(apical surface).
○ Polarity: One end attached to the basal layer, the other facing the lumen or
exterior.
● Types of Epithelium:
○ Covering and Lining Epithelium:
■ Simple: Single layer of cells (e.g., simple squamous, cuboidal, columnar).
■ Stratified: Multiple layers of cells (e.g., stratified squamous in skin,
mouth).
■ Pseudostratified: Appears layered but all cells rest on the basement
membrane (e.g., in the trachea).
 ■ Transitional: Specialized for stretching, found in the bladder.
○ Glandular Epithelium:
■ Glands: Specialized cells for secretion; categorized into:
■ Exocrine Glands: Have ducts (e.g., salivary glands).
■ Endocrine Glands: Secrete directly into the bloodstream.
■ Arise from epithelial germ layers (ectoderm and endoderm).
■ Can be multicellular or unicellular, with exocrine glands being tubular or
alveolar.
■ Myoepithelial Cells: Epithelial-derived, assist in secretion.

Connective Tissues

● Types: Bone, cartilage, fibrous connective tissue, adipose tissue, and blood.
● Functions: Support, storage, transport, and packing of organs.
● Structure: Composed of distinctive cells surrounded by an extracellular matrix (ECM),
which varies in consistency (liquid, gel-like, or hard).

Categories of Connective Tissue

1. General Connective Tissues:

○ Fibrous Connective Tissue:
■ Forms tendons and ligaments.
■ Main cell type: Fibroblast.
■ ECM: Network of protein fibers in a polysaccharide gel.
○ Mesenchyme:
■ An embryonic tissue, not a germ layer.
■ Non-polarized cells that can differentiate into various connective tissues.
2. Special Connective Tissues:
○ Bone:
■ Hard tissue with a matrix rich in calcium phosphate.
■ Cell types: Osteoblasts (build bone), Osteoclasts (break down bone),
Osteocytes (maintain bone).
■ Types: Cancellous (spongy) and compact bone.
■ Development: Endochondral (replacement of cartilage) and
intramembranous (direct formation from mesenchyme).
○ Cartilage:
■ Firm yet flexible tissue with a matrix of chondroitin sulfate and
collagen/elastic fibers.
■ Cell type: Chondrocytes in lacunae.
 ■ Types:
■ Hyaline Cartilage: Most common; found in joints, ribs, and
respiratory tract.
■ Fibrocartilage: Reinforced with collagen; found in intervertebral
disks and joints.
■ Elastic Cartilage: Flexible; found in the ear and epiglottis.
○ Blood:
■ Liquid connective tissue; matrix is plasma.
■ Functions in transport of gases, nutrients, and waste.
○ Hemopoietic Tissues:
■ Form blood cells; includes myeloid (in bone) and lymphoid (in lymphatic
organs) tissues.

Unique Processes

● Mineralization: Inorganic ions deposited in the matrix (e.g., calcification in bone and
cartilage).
● Ossification: Specific to vertebrates; deposition of hydroxyapatite in the organic matrix
of bone.

Key Differences Between Bone and Cartilage

● Matrix Composition: Cartilage has chondroitin sulfate; bone has calcium phosphate.
● Vascularization: Cartilage is avascular (no direct blood supply); bone is vascular.
● Microarchitecture: Bone is organized into osteons; cartilage is less organized.

Bone Development and Growth

Types of Bone Development

1. Endochondral Bone Development:

○ Definition: Bone forms from a cartilage model.
○ Structure:
■ Diaphysis: Middle shaft of the bone.
■ Epiphysis: Ends of the bone.
■ Metaphysis: Growth plate between diaphysis and epiphysis.
○ Process:
■ Mesenchymal cells condense to form a hyaline cartilage model.
■ A periosteal bone collar forms around the diaphysis.
■ Cartilage in the center calcifies, leading to chondrocyte death.
 ■ Blood vessels invade, creating a marrow cavity.
■ Ossification occurs as osteoblasts replace calcified cartilage with bone.
■ Growth continues through the epiphyseal plate, crucial for lengthening
bones.
■ Secondary ossification centers form in the epiphyses, further contributing
to bone growth.
2. Intramembranous Bone Development:
○ Definition: Bone forms directly from mesenchyme without a cartilage precursor.
○ Process:
■ Mesenchyme compacts into sheets, becoming highly vascularized.
■ Dense bars of bone matrix are deposited, with osteoblasts appearing.
■ Growth occurs by adding layers of new bone onto existing bars.
○ Types of Bones:
■ Dermal Bone: Formed within the skin's dermis (e.g., skull bones).
■ Sesamoid Bone: Forms within tendons (e.g., patella).
■ Perichondral/Periosteal Bone: Develops from connective tissue layers
covering cartilage or bone.

Bone Histology

● Bone is often organized into osteons, particularly in gnathostomes.

● In teleost fishes, bone can be acellular.
● Amphibians and reptiles typically have lamellar and cellular bone.
● In many mammals, large areas of bone may lack osteons.

Bone Remodeling and Repair

● Continuous Process: Bone remodels throughout life, adapting to mechanical demands.

● Repair Process:
1. Blood Clot Formation: Seals broken ends.
2. Callus Development: Forms from periosteum activity, comprising cartilage and
bits of the blood clot.
3. Replacement by Bone: The cartilaginous callus is gradually replaced by bone,
similar to endochondral ossification.
4. Final Remodeling: Osteoclasts and osteoblasts refine the repair, which can take
months to years.
Joints

Joints, or articulations, form where separate bone or cartilage elements come into contact. They
can be classified both functionally and structurally.

Functional Classification

● Synovial Joints (Diarthroses): Allow considerable movement.

● Synarthroses: Restrictive joints that permit little to no movement.

Structural Classification

● Synovial Joints: Defined by a synovial capsule made of dense fibrous connective tissue,
lined by a synovial membrane that secretes lubricating synovial fluid. The ends of the
bones are capped with articular cartilage.
● Synarthroses: Lack synovial structures and are further categorized:
○ Synostosis: Fusion of bones.
○ Synchondrosis: Connection via cartilage.
○ Syndesmosis: Connection via fibrous connective tissue.

Examples

● Symphyses: Cartilaginous joints, such as the mandibular and pubic symphyses.

● Sutures: Typically syndesmoses found in the skull.

Variability Across Species

While the dual classification system is primarily based on mammalian joints, variations exist in
other vertebrates:

● In snakes, the mandibular symphysis allows significant movement.

● In dolphins, individual phalanges may exhibit syndesmoses for structural strength.
● In birds, cranial bones may allow flexion despite being structurally syndesmoses,
facilitating cranial kinesis.

Neural Crest and Ectodermal Placodes

Neural crest cells and ectodermal placodes play crucial roles in vertebrate development and
evolution. While known since the nineteenth century, their significance has garnered more
attention in recent research.
Neural Crest Cells

● Origin: Neural crest cells are set aside early in vertebrate development. As the neural
folds form, these cells detach from the surface epithelium and gather into distinct cords
above the developing neural tube.
● Migration: Once released, they migrate along defined pathways within the embryo to
specific sites where they differentiate into various structures.

Derivatives of Neural Crest Cells:

● Ganglia: Form the spinal and cranial nerve ganglia.

● Schwann Cells: Create the insulating myelin sheath around peripheral nerves.
● Chromaffin Cells: Produce hormones in the adrenal medulla.
● Pigment Cells: Develop into various body pigment cells (excluding the retina and CNS).
● Connective Tissues: Contribute to cartilage and bone of the lower jaw and connective
tissue of voluntary muscles.
● Odontoblasts: Generate the inner layer of dentin in teeth.

These derivatives underline the extensive influence of neural crest cells in vertebrate anatomy
and function.

Ectodermal Placodes

● Definition: Ectodermal placodes are distinct from neural crest cells but may arise from
similar developmental processes. They typically appear as thickened areas of surface
ectoderm that sink inward to form sensory structures.

Types of Ectodermal Placodes:

● Otic Placodes: Develop into structures for hearing and balance (vestibular apparatus).
● Olfactory Placodes: Form at the head’s tip and develop into olfactory sensory receptors
connected to the brain.
● Adenohypophyseal Placode: An unpaired placode that contributes to the pituitary gland
(Rathke’s pouch).
● Optic Placodes: Lateral placodes that develop into the lens of the eye.

Interaction and Development

● Cranial and spinal nerves originate from both neural crest cells and ectodermal placodes,
indicating their integrated roles in sensory and nervous system development.
● In fishes and amphibians, placodes associated with cranial nerves are organized into two
rows, with upper dorsolateral and lower epibranchial placodes near the gill slits.
 ● Dorsolateral placodes also contribute to the lateral line system, while the otic placode is
crucial for auditory and vestibular function.

Significance of Neural Crest Cells and Ectodermal Placodes

● Wide Range of Structures: Contribute to many critical structures, especially in the head
region of vertebrates.
● Distinctive Features: Unique derivatives help distinguish vertebrates from other
chordates.
● Evolutionary Importance: Highlight the significance of these developmental processes
in vertebrate evolution.
● Integration: Derivatives from both neural crest cells and placodes integrate
harmoniously in adult vertebrates.
● Complexity and Adaptability: Reflect the complexity and adaptability of vertebrate
anatomy.

Extraembryonic Membranes

● Definition: Membranes that arise from embryonic germ layers, surrounding the
developing embryo.
● Function:
○ Sequester waste products.
○ Transport nutrients.
○ Exchange respiratory gases.
○ Create a fluid-filled capsule for the embryo.
○ Protect the embryo in a moist environment, reducing reliance on external water.
● Classification:
○ Amniotes: Reptiles, birds, and mammals with extraembryonic membranes (e.g.,
amnion).
○ Anamniotes: Fishes and amphibians, which lack most extraembryonic
membranes but have yolk sacs.

Development in Reptiles and Birds

● Formation: Extraembryonic membranes form early after germ layer establishment.

● Types:
○ Yolk Sac: Formed from splanchnopleure (endoderm + splanchnic mesoderm);
vascular network for nutrient mobilization.
 ○ Amnion: Formed from somatopleure (surface ectoderm + somatic mesoderm);
surrounds the embryo in an amniotic cavity.
○ Chorion: Peripheral layer formed from the somatopleure.
○ Allantois: Grows from hindgut endoderm; involved in waste storage and gas
exchange.

Development in Mammals

● Homologous Structures: Extraembryonic membranes are similar to those in reptiles and

birds.
○ Monotremes have membranes formed similarly to reptiles and birds.
○ Marsupials and eutherians have distinct processes for amnion and yolk sac
formation.
○ Chorioallantoic Membrane: Fusion of allantois and chorion; important for
respiration and nutrient exchange.

Eutherian Placenta

● Definition: Composite structure formed from both fetal and maternal tissues, allowing
vascular contact.
● Types:
○ Chorioallantoic Placenta: Formed from the chorioallantoic membrane; supplies
blood vessels from the allantois.
○ Yolk Sac Placenta: Formed from the yolk sac; provides nutrient exchange.
● Function:
○ Supports fetal respiration and nutrition.
○ Hormones produced help maintain the uterine environment.

Other Placentae

● Occurrence: Placentae are found in marsupials and some fishes, amphibians, and
reptiles.
● Types in Marsupials:
○ Yolk Sac Placenta: Common; facilitates gas and nutrient exchange.
○ Allantoic Placenta: Present in some species.
● Reptilian Placentae: Live-bearing reptiles have yolk sac and allantoic placentae, with
structures like placentomes for nutrient and gas exchange.
Overview of Early Embryonic Development

● Yolk Influence:
○ Yolk stores impact cleavage patterns and gastrulation.
○ Large yolk accumulations restrict cleavage to the yolk-free animal pole.
○ In teleosts, reptiles, birds, and monotremes, cleavage is discoidal (blastodisc
forms a cap on the yolk).
● Gastrulation:
○ Involves rearrangement of surface cells through embryonic structures (embryonic
shield or primitive streak).
○ These structures act as organizing areas, possibly homologous to blastopores but
adapted for large yolk.
● Evolution of Cleavage Patterns:
○ Discoidal cleavage evolved independently in teleosts and in
reptiles/birds/monotremes.
○ Modern amphibians show mesolecithal eggs with holoblastic cleavage, suggesting
a derived condition for discoidal cleavage in amniotes.
○ Eutherian mammals show discoidal cleavage despite minimal yolk, reflecting
ancestral traits.
● Amniotes vs. Anamniotes:
○ Division into amniotes (with extraembryonic membranes) and anamniotes
(without) reflects differences in embryonic support.
○ Amnion and other membranes are adaptations for terrestrial life, allowing the
development of shelled, cleidoic eggs.

Development of the Coelom and Its Compartments

● Coelom Formation:
○ Originates from the hypomere during early development.
○ Subdivided into anterior pericardial cavity (heart) and posterior pleuroperitoneal
cavity (viscera).
● Transverse Septum:
○ Fibrous partition separating the coelomic compartments.
○ Embryonic veins pass through, connecting with the developing liver.
● Pleural Cavities in Reptiles and Birds:
○ Some reptiles (e.g., crocodiles, turtles) and all birds have pleural cavities.
○ Pulmonary fold (nonmuscular septum) isolates lungs in pleural cavities.
● Diaphragm Development in Mammals:
○ Coelomic fold (pleuroperitoneal membrane) fuses with transverse septum,
forming a muscular diaphragm.
 ○ Diaphragm contributes to lung ventilation; muscle origins are complex, involving
thoracic and cervical myotomes.
● Mesothelium:
○ Lines the coelom, providing structural integrity and compartmentalization.
○ Facilitates organ movement and prevents conflicting activities (e.g., separates
heart from viscera).
● Functionality of Coelomic Compartments:
○ Allows localized control over internal organs.
○ Mesenteries (connective tissue structures) suspend organs and provide pathways
for blood vessels and nerves.
● Scrotum in Mammals:
○ A coelomic pouch for testicular descent, creating a cooler environment for sperm
production.

Maturation and Metamorphosis

● Metamorphosis:
○ Transition from larval to adult form in organisms where larvae are
morphologically different from adults.
○ Metamorphosis can be radical and involves significant structural changes.
○ In vertebrates, maturation occurs from juvenile to adult, and ontogeny continues
throughout life, not ending at hatching or birth.
● Larval Stage:
○ Larvae are often mobile and may inhabit different environments from adults,
serving as dispersal stages (e.g., tunicate larvae).
○ Tadpoles (frog larvae) utilize transient resources and are more limited to aquatic
habitats compared to adult frogs.
○ Variability in duration and form of larval stages can be adaptive; for example,
lampreys may remain larval for years if conditions favor it.
● Adaptations in Development:
○ Some species (like certain salamanders) can forgo metamorphosis altogether,
becoming sexually mature while retaining larval characteristics.
○ Conditions may favor direct development, where embryos develop directly into
adult forms, bypassing larval stages to avoid predation (e.g., Puerto Rican tree
frogs).

Heterochrony

● Definition:
 ○ Heterochrony refers to changes in the timing of developmental events compared
to ancestral species, influencing evolutionary trajectories.
● Types of Heterochrony:
○ Paedomorphosis: Retention of juvenile features in adult forms of descendants
(e.g., mudpuppies retain gills).
○ Peramorphosis: Exaggeration of ancestral adult features in descendants (e.g.,
elongated limbs).

Mechanisms of Heterochrony

● Peramorphosis Processes:
○ Hypermorphosis: Prolonged development leads to larger adult forms.
○ Acceleration: Features grow faster during ontogeny compared to ancestors.
○ Predisplacement: Features develop earlier in the life cycle, giving them a
developmental head start.
● Paedomorphosis Processes:
○ Progenesis: Early cessation of growth leads to juvenile characteristics in adults.
○ Neoteny: Slower growth allows juveniles to reach sexual maturity before
completing development.
○ Postdisplacement: Late appearance of features results in retention of juvenile
traits.

Ecological and Evolutionary Implications

● The balance of time spent in larval versus adult forms can shift based on environmental
conditions.
● Evolutionary changes in timing can produce rapid adaptations and new morphologies.
● The entire ontogeny of an individual is an adaptive response to its environment,
influencing evolutionary pathways.

Recapitulation

● Definition and Debate:

○ Recapitulation suggests that juvenile forms of descendant species resemble adult
forms of ancestors.
○ Historically controversial, often tied to the "biogenetic law" (ontogeny
recapitulates phylogeny).
○ Modern perspectives question its validity, preferring clearer terminology related
to heterochrony.
Ontogeny and Phylogeny

Biogenetic Law

● Concept Origin: Proposed by Ernst Haeckel in 1866, stating that ontogeny (individual
development) reflects phylogeny (evolutionary history).
● Embryonic Evidence: Features such as pharyngeal slits appear in the early embryos of
reptiles, birds, and mammals, suggesting a common ancestry with fish, even if these
features are lost before birth.
● Recapitulation: Haeckel suggested that the development of an individual is a condensed
version of its evolutionary past, although he acknowledged some exceptions.

Critiques and Alternatives

● Karl Ernst von Baer’s Counterargument: Von Baer argued that development moves from
general to specific, meaning that embryos of descendant species diverge from their
ancestors rather than resembling them. For example, chick embryos do not show features
of adult fish.
● General to Specific Development: Development begins with undifferentiated cells,
progressing through layers and tissues to form specific organs, leading to unique features
that do not mirror ancestral adults.

Overview of the Biogenetic Laws

● Misapplication of Haeckel's Law: There is no direct correspondence between

descendant embryos and the adults of ancestors. Instead, embryos resemble their
ancestors at early developmental stages.
● Common Patterns: It is common for embryos to display similar structures initially,
followed by differentiation into species-specific traits (e.g., limb buds becoming fins,
wings, or limbs).

Conservatism in Development

● Embryonic Features: Certain features, like gill slits, may persist during development but
do not necessarily have functional roles in adults, reflecting a kind of developmental
conservatism rather than true recapitulation.
● Functional Roles: Structures like the notochord in embryos serve critical roles in
development, laying the groundwork for later structures such as the vertebral column.

Reasons for Developmental Conservatism

 ● Pleiotropy: Genes can have multiple effects, complicating the modification of traits.
Altering one gene can disrupt several interconnected traits, leading to a conservative
nature in development.
● Evolutionary Remodeling: Innovations often arise through the modification of existing
structures rather than new constructions. For example, forelimbs may evolve into wings,
and adaptations like bipedalism in humans arise from reconfigurations of ancestral traits.

Hox Genes and Their Kingdoms

Homeotic Genes

● Definition: Homeotic genes, a term coined by William Bateson, refer to genes that control
the development of body structures in animals and plants. Mutations can cause parts of an
organism to transform into other parts.
● Examples: In fruit flies, homeotic mutations can lead to legs replacing antennae or an
extra pair of wings forming.

The Role of Hox Genes

● Master Control Genes: Hox genes act as regulatory genes that govern the expression of
numerous structural genes responsible for body part formation. They play a critical role
in establishing the body plan during development.
● Conservation Across Species: Hox genes are found throughout the animal kingdom,
including in plants and fungi, highlighting their evolutionary significance.

From Egg to Adult

● Development Process: The journey from a single-celled egg to a complex adult

organism involves numerous cell divisions. Initially, these divisions are limited to
cleaving the egg but later contribute to the embryo's growth.
● Cellular Equivalence: Although all cells in the embryo carry the same DNA, their
functions are directed by the activation of specific genes at the right time and place.

Establishing Body Polarity

● Body Symmetry: Early embryonic development establishes body polarity, defining

anterior (front), posterior (back), dorsal (top), and ventral (bottom) regions. This is often
achieved through chemical gradients that guide the formation of body structures.
● Hox Gene Activation: In some organisms, Hox genes are involved in setting up this
body polarity, helping to position body parts accurately.
Building the Embryo

● Activation of Hox Genes: Once body polarity is established, Hox genes are activated by
positional and environmental cues, influencing the expression of structural genes
necessary for forming body parts.
● Master Control Function: A single Hox gene can regulate the expression of many
structural genes, making small changes in Hox genes potentially result in significant
morphological changes.

Evolutionary Significance of Hox Genes

● Gene Duplication: Research shows that duplications of Hox genes and increases in their
numbers correlate with major evolutionary transitions among animal phyla.
● Segment Development: Changes in the number of body regions governed by Hox genes
can lead to the addition or alteration of body segments.

Complexity and Adaptation

● Conservation and Variation: Hox genes are highly conserved in their sequences and
expression patterns across different species. However, the mechanisms of activation and
the responses of downstream genes can evolve.
● Dynamic Role in Development: Some Hox genes are activated multiple times during
development, leading to different effects based on the timing and context of their
activation. For example, halteres in flies are modified wing structures that demonstrate
evolutionary adaptation.

Epigenomics: Overview

Beyond Just Genes

Organisms are shaped by more than just their genes. While genes produce RNA that eventually
makes proteins, the actual development of an organism relies heavily on interactions between
cells and tissues. This field of study is called epigenomics, which looks at how genes are
regulated without changing the DNA itself.

Induction in Development

● Sclerotomes and Neural Tube: Early in development, groups of tissues called

sclerotomes form next to the neural tube, which develops into the spinal cord. If part of
the neural tube is removed, not only does the spinal cord fail to develop, but the nearby
 sclerotomes also don’t form properly. This process is known as induction, where one
tissue influences the development of another.
● Limbs Development: In developing limbs, the apical ectodermal ridge (AER) and the
underlying mesoderm work together. The AER encourages limb growth, while the
mesoderm decides whether the limb will be a forelimb or hindlimb. If the AER is
removed, limb development stops, showing how important these interactions are.

Evolutionary Implications

● Adaptation: Changes in how tissues interact can lead to significant evolutionary

adaptations. For example, some lizards have lost their legs to better move through their
environments. This change happens because alterations in the interaction between tissue
blocks prevent limb formation.
● Hox Genes: These are master control genes that help set up body plans. In limbless
snakes, certain Hox genes don’t activate the AER, leading to the loss of limbs. This
shows how small changes in development can result in major evolutionary shifts.

Changes in Birds and Horses

● Birds: The evolution of birds shows changes in the bones of their legs. Experiments can
sometimes recreate traits of their ancestors, suggesting that minor developmental changes
could have led to significant differences over time.
● Horses: Modern horses have just one toe on each leg, unlike their multi-toed ancestors.
This reduction likely happened through changes in developmental patterns,
demonstrating that major adaptations can arise from a few genetic changes.

Overview of Organism Development

Organisms go through a life history that includes several stages: starting as a fertilized egg,
undergoing embryonic development, maturing (often through a larval stage), reaching sexual
maturity, and eventually aging before death.

Key Stages of Development

1. Differentiation: As the embryo develops, cells become specialized through a process

called differentiation. This leads to diverse cell types that will form organs and body
structures.
2. Morphogenesis: This is the process by which cells move and organize to create the
body’s shape. Before fertilization, the egg may have uneven distributions of materials,
which help define the future body layout.
 3. Cleavage and Blastula Formation: After fertilization, the single-celled zygote
undergoes rapid division (cleavage), forming a hollow ball of cells known as a blastula.
4. Gastrulation and Neurulation: During these phases, three primary germ layers
(ectoderm, mesoderm, endoderm) form through processes like:
○ Epiboly: Spreading of cell layers
○ Involution: Folding of layers inward
○ Invagination: Infolding to create pockets
○ Delamination: Splitting into layers
5. The neural tube, which becomes the spinal cord, forms through a process called primary
neurulation.
6. Germ Layers and Tissue Formation: The three germ layers give rise to four basic
tissue types: epithelial, connective, muscle, and nervous tissues. These tissues work
together to form organs.
7. Extraembryonic Membranes: In amniotes (like reptiles and mammals), not all cells
from the fertilized egg contribute to the embryo. Some form membranes that support the
embryo's development, providing nutrition and protection.

Maturation and Sexual Maturity

After birth or hatching, the organism may spend time as a larva or juvenile before reaching
sexual maturity. Maturation can happen gradually or suddenly (metamorphosis).

Evolutionary Changes in Development

Heterochrony refers to changes in the timing of developmental events, which can lead to
significant evolutionary changes. These changes can occur in:

● Onset: When a part begins to grow

● Rate: How quickly it grows
● Offset: When growth stops

These timing alterations can lead to variations such as:

● Peramorphosis: Overdevelopment of features

● Paedomorphosis: Retaining juvenile traits into adulthood

Conservatism and Variation

While there is a lot of variation in how embryos develop, especially due to yolk content, some
processes remain consistent across different vertebrate groups. For example, eutherian mammals
(which have less yolk) still show patterns similar to those with more yolk, indicating a
conservative nature in developmental processes.
Genetic and Developmental Connections

Common genetic and tissue processes are fundamental to development. Key points include:

● Patterning: Early embryonic stages establish basic body layouts (dorsal, ventral,
anterior, posterior).
● Cell Signaling: Cells communicate and coordinate actions through signaling, either by
direct contact or through signaling molecules.

Hox Genes: These act as master control switches for development, regulating when and where
other genes are activated. They do not directly code for proteins but influence overall body plan.

Conclusion

Development is a complex interplay of genetic regulation and tissue interactions. There’s no

simple one-to-one relationship between genes and body parts; instead, small genetic changes can
lead to significant structural modifications. This understanding emphasizes how interconnected
all parts of an organism are and how evolution can bring about rapid and substantial changes
through subtle genetic shifts.