Chapter 6: Integumentary System

The integument, or skin, is a complex organ with multiple layers and functions. It consists of:

1. Epidermis: The outer layer, derived from the ectoderm.

2. Dermis: The inner layer, developed from mesoderm.
3. Basement Membrane: A thin layer between the epidermis and dermis, composed of
basal and reticular lamina.

Beneath the dermis lies the hypodermis (or superficial fascia), made of loose connective and
adipose tissues.

Importance of the Integument

The integument is one of the largest organs, accounting for about 15% of body weight in
humans. It produces a variety of structures, including:

● Hair
● Feathers
● Claws and nails
● Horns and beaks
● Scales

Both the epidermis and dermis work together to form structures like teeth and scales. If one is
missing, the other can't form these specialized features effectively.

Functions of the Integument

The skin serves many essential roles:

● Barrier Protection: It protects against pathogens and physical injury.

● Temperature Regulation: It helps maintain body heat and releases excess heat.
● Sensory Perception: Contains receptors for touch, pain, and temperature.
● Osmoregulation: Aids in regulating water and ion movement.
● Coloration: Skin pigments protect against UV light and can play a role in mating
displays.

Embryonic Development of the Integument

By the end of neurulation, skin precursors are largely formed:

 ● The ectoderm thickens to create the multilayered epidermis, with the deepest layer
(stratum basale) resting on the basement membrane. This layer continuously produces
new cells to replace the outer layer (periderm).

The dermis originates from:

● The dermatome, a part of the somite that develops into the dermis' connective tissue.
● Cells from the neural crest contribute to pigment cells (chromatophores) and bony
structures in the skin.

Structural Composition

The integument consists of two primary layers—epidermis and dermis—along with the basement
membrane, blood vessels, and nerves. These elements allow for a wide variety of skin features.

Skin Glands

The epidermis can invaginate to form skin glands:

● Exocrine glands: Retain ducts to release substances to the skin surface.

● Endocrine glands: Lose ducts and release hormones directly into the bloodstream.

General Features of the Integument

Dermis

The dermis in many vertebrates is responsible for producing dermal bones through a process
known as intramembranous ossification. These bones, found prominently in ostracoderm
fishes, also appear in some mammals. The primary component of the dermis is fibrous
connective tissue, mainly composed of collagen fibers, which can be organized into distinct
layers called plies.

● Collagen Arrangement:
○ In species like the amphioxus, collagen fibers are arranged in a highly ordered
manner, forming plies that create a supportive structure. This organization helps
prevent sagging and provides shape to the skin.
○ In aquatic vertebrates (like sharks), collagen bundles are oriented at angles,
allowing for flexibility and resilience during movement. This arrangement enables
the skin to stretch without wrinkling, ensuring smooth water flow across the body.
● Stratum Compactum:
 ○ In fish and some aquatic vertebrates, this layer is well-defined, whereas it is less
pronounced in terrestrial vertebrates, where limb motion is more crucial.

Epidermis

The epidermis serves several important functions across vertebrate groups:

● Mucus Production:
○ In many fish, the epidermis secretes mucus, which helps protect against bacterial
infections and promotes laminar water flow over the body. In amphibians, mucus
prevents drying during terrestrial life.
● Keratinization:
○ Terrestrial vertebrates have a keratinized layer known as the stratum corneum,
composed entirely of dead cells that reduce water loss in dry environments.
○ New cells arise in the stratum basale, pushing older cells upward, where they
undergo keratinization, forming a protective outer layer.
○ There are two types of keratinocytes: alpha-keratin (soft, flexible) and
beta-keratin (hard, found in scales, claws, etc.).
● Scales Formation:
○ Scales arise as folds in the integument. If dermal contributions are dominant, they
are termed dermal scales; if epidermal, they are epidermal scales.

Phylogeny of the Integument

Integument of Fishes

Most living fishes have nonkeratinized skin covered with mucus, which helps with protection
and flow dynamics in water.

● Epidermis Composition:
○ Contains epidermal cells and specialized unicellular glands, including club cells
(for alarm signaling) and goblet cells (for mucus production).
● Dermis:
○ The dermis consists of regular layers of collagen, allowing flexibility without
wrinkling, and gives rise to dermal bone and scales.

Primitive Fishes

● Ostracoderms and Placoderms:

 ○ These fish had bony dermal plates providing significant protection. The surface
often featured tubercles made of enamel-like substances over a dentin layer.
● Lampreys and Hagfishes:
○ Lacking dermal bone, their skin is smooth and covered in living cells with
specialized glands.

Chondrichthyes (Cartilaginous Fishes)

● They lack dermal bone but have placoid scales, which enhance hydrodynamics during
swimming. The dermis has a layered fibrous structure providing strength.

Bony Fishes

● Their dermis is divided into a superficial layer (loose connective tissue) and a deeper
layer (dense fibrous tissue).
● Scales:
○ Types include cosmoid scales (thick dentin layer), ganoid scales
(enamel-coated), and teleost scales (lacking enamel and dentin).
○ Teleost scales can be further classified into cycloid (smooth) and ctenoid
(fringed) types. These scales exhibit growth rings that can be used to determine
the age of the fish.

Integument of Tetrapods

General Features

● Keratinization: Extensive in tetrapods, forming the protective stratum corneum.

● Function: Resists abrasion and prevents moisture loss.
● Gland Types: Multicellular glands in tetrapods; fish have unicellular mucous glands.

Amphibians

● Metamorphosis: Transition from aquatic larvae to terrestrial adults.

● Skin Function: Serves as a respiratory surface for gas exchange.
● Skin Structure:
○ Aquatic larvae: Simple dermis; epidermis with large Leydig cells for protection.
○ Terrestrial adults: Defined layers (basale, spinosum, granulosum, corneum).
● Glands: Mucous and poison glands; nuptial pads form during breeding.

Reptiles
 ● Adaptations: More extensive keratinization; fewer glands.
● Scales: Epidermal scales, lacking bony support from dermis.
● Skin Layers: Includes stratum basale, granulosum, and corneum.
● Molting: Different patterns; some shed large pieces (e.g., snakes).

Birds

● Feathers: Unique keratinized structures, essential for flight.

● Skin Features:
○ Rich vascularization around feather follicles.
○ Few glands; uropygial gland for feather maintenance.
● Feather Types:
○ Contour Feathers: Shape body.
○ Down Feathers: Provide insulation.
○ Flight Feathers: Long, aerodynamic, asymmetrical vanes.

Evolution of Feathers

● Initial Function: Likely for insulation before adaptation for flight.

● Modern Structure: Features like barbs, barbules, and rachis enhance flight efficiency.
● Historical Evidence: Early birds (e.g., Archaeopteryx) had features suitable for flight.

Integument of Mammals

General Structure

● Layers of Skin:
○ Epidermis: Outermost layer, specialized as hair, nails, or glands.
○ Dermis: Underlying layer with connective tissue and fat (hypodermis).

Epidermis

● Keratinocytes: Main cell type, forms the protective, cornified layer.

● Layers:
○ Stratum Basale: Deepest layer; cells divide and push outward.
○ Stratum Spinosum: Provides strength.
○ Stratum Granulosum: Contains keratin.
○ Stratum Lucidum: Present in thick skin (e.g., soles).
○ Stratum Corneum: Outermost layer, continuously exfoliated.
● Other Cell Types:
○ Langerhans Cells: Immune function.
 ○ Merkel Cells: Mechanoreceptors for touch.
○ Chromatophores: Produce melanin, affecting skin color.

Dermis

● Layers:
○ Papillary Layer: Contains dermal papillae; supports epidermis.
○ Reticular Layer: Dense, irregular connective tissue; houses blood vessels,
nerves, and hair follicles.
● Function: Anchors skin to underlying structures, supports hair follicles and glands.

Hair

● Structure:
○ Root: Embedded in the follicle.
○ Shaft: Visible part, composed of cuticle, cortex, and medulla.
● Growth Cycle:
○ Growth Stage: Active cell proliferation.
○ Degeneration Stage: Inactive cells.
○ Rest Stage: Follicle inactive until new growth begins.
● Color: Determined by pigments from chromatophores.

Types of Hair

● Guard Hairs: Coarse, outer layer.

● Underfur: Finer, insulating layer.
● Vibrissae: Whiskers with sensory function.

Glands

● Sebaceous Glands: Produce sebum; condition and waterproof skin.

● Eccrine Glands: Produce watery sweat; not associated with hair follicles; involved in
thermoregulation.
● Apocrine Glands: Viscous secretions; associated with hair follicles; involved in scent
marking and signaling.
● Mammary Glands: Produce milk; consist of lobules with secretory alveoli; functional
primarily in females.

Mammary Glands

● Structure: Lobules, alveolar ducts, and teats/nipples.

● Function: Lactation; nourishes young.
● Evolution: Hypothesized to have evolved from skin gland secretions that initially
protected and then nourished offspring.

Specializations of the Integument

Nails, Claws, and Hooves

● Nails:
○ Found only in primates.
○ Protect digit tips; provide stability for grasping.
○ Form from the nail matrix at the base.
● Claws:
○ Curved, laterally compressed structures.
○ Common in amphibians, birds, reptiles, and mammals.
● Hooves:
○ Found in ungulates (e.g., horses).
○ Structure includes hoof wall, sole, and frog.
○ Absorb impact and support weight.
○ Grow from the germinal region (matrix cells) at about 6 mm per month.

Horns and Antlers

● Horns:
○ Found in Bovidae (e.g., cattle, goats).
○ Unbranched, keratinized sheath over a bony core.
○ Retained year-round; grow continuously.
● Antlers:
○ Found in Cervidae (e.g., deer).
○ Branched, shed annually; covered in velvet during growth.
○ Hormonal control for growth and shedding.

Baleen

● Keratinized plates in mysticete whales.

● Function as strainers to extract krill from water.
● Formed from the integument, not bone.

Scales

● Serve protective and functional roles:

○ Prevent mechanical damage and pathogen entry.
 ○ Minimize water loss.
○ Enhance locomotion in water (e.g., in fishes).
● Found in reptiles, some mammals (e.g., beaver).

Dermal Armor

● Formed from dermal bone in some fish (e.g., ostracoderm, placoderm).

● Contributes to turtle shells (carapace and plastron).
● Represents evolutionary link between integument and skeleton.

Mucus

● Inhibits pathogens; aids in moisture retention for gas exchange in amphibians.

● Reduces friction during swimming in aquatic species.

Coloration

● Results from:
○ Physical properties: Light scattering (e.g., Tyndall scattering).
○ Chemical properties: Pigments in chromatophores.
● Types of chromatophores:
○ Melanophores: Contain melanin; protect against UV and add color.
○ Iridophores: Reflect light; cause iridescence.
○ Xanthophores: Contain yellow pigments.
○ Erythrophores: Contain red pigments.
● Color Change Mechanism:
○ Chromatophores assume a relatively fixed shape.
○ Hormonal stimulation causes pigment granules to be shuttled into pseudopods or
concentrated centrally within the cell.
● Color changes can occur for camouflage or due to hormonal responses.

Overall Overview of the Integumentary System

● Skin as a Dynamic Organ:

○ The epidermis is replaced approximately every two weeks.
○ Stem cells in the epidermis are responsible for this turnover.
● Homology of Integumentary Structures:
○ Hair, feathers, and reptilian scales are products of the epidermis and are broadly
homologous.
○ Controversies exist regarding their specific evolutionary origins:
 ■ Some propose hair is a transformed scale; others argue it derives from
sensory bristles.
■ Similarities between placoid scales and teeth lead to debates on their
evolutionary relationship.
● Layers of Skin:
○ Skin consists of two main layers: the epidermis and the dermis.
○ The dermis regulates and influences epidermal cell proliferation and
differentiation.
● Experimental Embryology Insights:
○ Isolated epidermis can survive but doesn’t proliferate without dermis.
○ Epidermis from a chick leg can form scales when combined with embryonic
dermis; fails with other tissues.
○ Epidermis shows some autonomy by forming a keratinized layer when exposed to
air.
● Influence of Dermis on Epidermis:
○ Dermis directs epidermal differentiation (e.g., scales vs. feathers).
○ Transplanting dermis affects the type of epidermal derivatives formed.
○ In species like lizards, dermis type determines scale shape and type.
● Inductive Interactions:
○ Dermis does not directly contribute cells but induces epidermal specializations.
○ Cross-species transplants show epidermis typically retains its characteristic
specialization.
○ Transplants may induce some epidermal proliferation but can fail to fully develop
typical structures.
● Evolution of Skin:
○ Evolutionary changes involve dermal induction capacity and epidermal response.
○ The interaction between epidermis and dermis is crucial for normal skin
derivative formation.
○ Notable experiments show potential for tooth formation in birds when combined
with mouse dermis.
● Conclusion:
○ The interactions between epidermis and dermis represent an evolving unit
influencing skin specialization.
○ Broad homology exists among integumentary structures, including hair, feathers,
scales, teeth, and bony-fish scales.