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Structure and function of the eye and its adnexa
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3D Ophthalmology Introduction Vision is the ability of living creatures to interpret their environment through rays of light (photons). The visual system comprises the eyes, the connecting pathways, and the visual cortex. In order for an animal to see, light photons must enter their eyes and be focused correctly on the retina. Retinal photopigments convert this light energy into electrical energy. All the properties of the image (brightness, orientation, depth, etc.) are subsequently interpreted by the visual cortex, which extracts the most relevant information. The eye, which is almost perfectly spherical, is one of the most highly specialised organs. The main function of the eye is to focus the image and regulate the amount of light that hits the retina, thereby reducing optical aberrations and ensuring the best possible visual acuity. Any change in the transparency or the physiological arrangement of the ocular structures can diminish visual capacity. The ocular adnexa encompass all of the eyeball’s supporting structures, that is, those surrounding the eye and which contribute to its function. These include, amongst others, the eyelids, conjunctiva, nasolacrimal system, and the nictitating membrane. The eyeball is located in the orbit, where it is held in place by the extraocular muscles and orbital fasciae. The adnexa and orbit are essential for the normal function of the eye.
Anatomy of the eye The eye is traditionally divided into three layers:
• The fibrous tunic: formed by the cornea and sclera. This is a collagen-rich layer that lends the eye its structure, and the anterior portion is transparent to allow light to enter the eye. The optic nerve emerges at the rear of the eye and passes through a structure called the cribriform plate. • The uvea: a highly vascularised layer, comprising the iris, ciliary body, and choroid. • The retina and optic nerve: the retina is responsible for transduction, that is, the act of transforming light into an electrical signal. The electrical information is then transmitted to the nervous system via the optic nerve. The crystalline lens, which helps focus the image on the retina, separates the anterior and posterior segments of the eye. The anterior segment is full of aqueous humour, which nourishes the structures contained within and helps
maintain intraocular pressure. Aqueous humour is secreted by the ciliary body and drains into the iridocorneal angle. The posterior segment contains vitreous humour, which is rich in collagen and helps shape the eye and hold the retina in place.
The cornea and sclera The cornea is the most anterior part of the fibrous tunic. It is a transparent layer which in dogs has a marginally greater width than height. It is around 0.5 mm thick. It features four layers: a stratified epithelium, the collagen stroma, Descemet’s membrane, and the endothelium. The epithelium contains basal, intermediate, and squamous cells which are all continuously renewed. It does not contain keratin, thus promoting transparency. The stroma makes up most of the cornea’s thickness. It is formed from collagen fibres and keratocytes, which are special fibroblasts charged with keeping the cornea transparent, healing ulcers, and synthesising the components of the stroma. Corneal collagen is organised in lamellae that are arranged orthogonally and at a fixed separation. This special configuration allows light to pass through and differentiates the cornea from other collagen-rich structures, such as the sclera, which are opaque. Furthermore, the stroma is rich in glycosaminoglycans, and again these also contribute to transparency. Some scarring processes can cause a loss of transparency in the cornea. Descemet’s membrane is the basement membrane for the corneal endothelium. It is an elastic layer that gets thicker as the animal ages. In the case of glaucoma, Descemet’s membrane can rupture, resulting in what are known as Haab’s striae. A corneal ulceration accompanied by a total loss of the stroma leads to exposure of Descemet’s membrane (descemetocele). The corneal endothelium is the most metabolically active layer. This monocellular layer dehydrates the cornea, thus helping to maintain its transparency. Corneal endothelium cells are postmitotic, so they are unable to regenerate, and their number decreases over the course of a lifetime. The remaining cells grow larger to fill the spaces left by those which die. If the number falls below a critical value, it becomes impossible to maintain a suitable balance between the amount of water that “enters” and “leaves” the stroma, leading to corneal decompensation and in turn oedema.
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3D Ophthalmology
Structure and function of the eye and its adnexa A healthy cornea should be smooth, shiny, avascular, and free from pigment. The corneal stroma is avascular and contains very few cells, so it receives oxygen and metabolites from tears, the aqueous humour, and sclera. Any alterations to these structures will cause changes in the cornea. A common example is dry eye, in which the cornea may accumulate pigment, experience neovascularisation, and lose its smooth surface. The cornea is one of the most sensitive parts of the body. It is innervated by the long ciliary nerves, which enter the anterior corneal stroma radially, and they are subsequently attached to the ophthalmic branch of the trigeminal nerve. Corneal sensitivity is greater at the surface and in the anterior stroma; therefore, as a lesion gets deeper, sensitivity declines. Brachycephalic dogs have less corneal sensitivity than mesocephalic or dolichocephalic breeds, which is why they show less signs of pain when affected by corneal ulcers.
The sclera makes up most of the fibrous tunic and is divided into three layers: the most superficial layer is called the episclera, the middle layer is the scleral stroma, and the deepest is the lamina fusca. The episclera provides the point of attachment between Tenon’s capsule and the scleral stroma. It is a highly vascularised fibrous layer. Like the cornea, the scleral stroma consists of collagen and fibroblasts;
Figure 1. Crystalline lens, iris, and ciliary body as seen from the retina.
however, in this case the collagen is disorganised, so the scleral stroma is not transparent. The lamina fusca is where the most external layers of the choroid and ciliary body attach to the sclera.
The uvea The uvea is the middle layer of the eye. It can be further divided into the anterior uvea, formed by the iris and ciliary body, and posterior uvea, comprising the choroid. The iris is a diaphragm that controls how much light enters the eye. This control is achieved with two muscles: the pupillary dilator and sphincter muscles. They open or close the pupil depending on how much light enters the eye, adjusting pupil size to ambient conditions. The anterior aspect of the iris does not have an epithelium and is composed of stromal cells. The posterior aspect has a pigmented epithelium. There are two arteries (long ciliary arteries), entering nasally and temporally, forming an arterial circle, which may or may not be complete. The ciliary body is located behind the iris (Fig. 1). The anterior section features a series of folds (pars plicata), known as ciliary processes. These folds subsequently become less prominent, until they give way to a flat area (pars plana) that eventually joins the retina at the ora serrata. The ciliary body is covered with an epithelial bilayer and has a muscle at its base formed from smooth muscle fibres innervated by the parasympathetic system. These fibres attach to the base of the ciliary body and are closely related to the iridocorneal angle. Uveitis causes the ciliary muscle to contract, which is painful and increases the amount of drainage via the normal route. The choroid can be found in the posterior uvea. It is a fine layer of very well vascularised tissue with a variable degree of pigmentation. Found uniquely at the back of the choroid is a triangular layer call the tapetum. Its function is to reflect light and therefore double the stimulation of the retina’s photoreceptor cells. The tapetum varies in size and colour, depending on the breed of dog, and may even be physiologically absent. One of the most important functions of the choroid is to act as the blood–ocular barrier. This separates the eye from the body’s general circulation to prevent proteins and macromolecules from filtering into the aqueous humour and vitreous humour. Inflammation can alter the barrier’s impermeability.
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3D Ophthalmology The crystalline lens The crystalline lens is a biconvex, avascular lens located inside the eye and suspended by ciliary zonular fibres. The anterior face is more planar than the posterior face and the lens has an equator, which is the circumference where the two faces meet.The crystalline lens is surrounded by a capsule and has an epithelium, which is located under the anterior capsule. The lens fibres are arranged in concentric layers, running out from the central nucleus and forming the cortex.These fibres are produced throughout the animal’s life, so they get pushed towards the lens nucleus. The main purpose of the lens is to accommodate for objects at different distances and focus the image on the retina. However, dogs have a very low capacity for accommodation.
The vitreous humour The vitreous humour is an elastic, transparent hydrogel that occupies a cavity with the same name and comprises up to 80 % of the eye’s volume. It mainly consists of water, collagen fibres, hyaluronic acid, and hyalocytes. Hyalocytes present phagocytic activity and can transform into fibroblasts and, therefore, form scar tissue. Anatomically, this gel can be divided into the anterior, posterior, peripheral, and central vitreous, and it is worth noting that the collagen fibres are especially dense in the peripheral area, which is where these fibres attach strongly to the surface of the retina. Degeneration and liquefaction of the peripheral vitreous can reduce the force it exerts on the retina, increasing its chance of detachment. There is another area where the vitreous fibres are firmly adhered to an intraocular structure. The crystalline lens sits in a depression in the vitreous called the hyaloid fossa, which is exactly the point where the posterior lens
capsule is strongly attached to the vitreous. Shifts of the lens, which accompany its luxation, produce traction in the vitreous fibres and, consequently, in the retina.
The retina and optic nerve The most important part of the eye is the retina. The ultimate objective of all the ocular structures is to focus an image on the photoreceptor cells and produce a picture with the greatest possible visual acuity. The retina is a very complex tissue composed of many different types of cell involved in generating the final electrical impulse. In simple terms, the retina contains photoreceptor cells that transmit the electrical impulse to the bipolar cells, which then transmit it to the ganglion cells. The axons of the ganglion cells form the optic nerve. Photoreceptors are comprised of cone and rod cells, both of which contain photopigments that react to light and produce an action potential.
Cone cells are sensitive to more intense light and provide colour vision, while rod cells are stimulated by less intense illumination.
The photoreceptors rely on the retinal pigment epithelium, as it performs several functions such as recycling photopigments. In the event of retinal detachment, the pigment epithelium always remains attached to the choroid. The optic nerve is formed when all the ganglion cell axons merge to establish the optic nerve head. The nerve then traverses the cribriform plate, through the cone formed by the extraocular muscles, and the optic foramen of the orbit, after which it joins with the optic nerve from the other eye at a point called the optic chiasm (Fig. 2).
Figure 2.
Eyeball with the departing optic nerve.
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3D Ophthalmology
Structure and function of the eye and its adnexa
Anatomy of the ocular adnexa Eyelids
The eyelids are musculocutaneous flaps vital to maintaining a healthy ocular surface. They comprise the tarsus (a fibrous plate that provides structure), muscles that control its position and contraction, elastic skin on the external surface, and palpebral conjunctiva on the internal surface. Dogs only have eyelashes on the upper eyelid (Fig. 3). There are some glands on the eyelid margin, called meibomian or tarsal glands, that secrete a mixture of lipids. The glands’ orifices can be examined with an optical magnification system. There are some other glands, e.g. the glands of Zeis and Moll, but they are clinically insignificant. Principally, the eyelids are closed by contracting the orbicularis oculi muscle, which is situated in the anterior portion of the tarsus of both eyelids. In dogs, the upper eyelid is more mobile than the lower eyelid and innervated by the oculomotor nerve, unlike other palpebral muscles which are innervated by the facial nerve. The eyelids have a ligament in the medial canthus and a muscle that serves as a ligament in the lateral canthus, the retractor muscle of the lateral angle (retractor anguli oculi lateralis). Sensory information from the eyelids is transmitted via the trigeminal nerve. When pain is detected, the orbicular muscle contracts strongly, resulting in blepharospasm. The eyelids protect the eyes and help distribute and drain tears. When blinking, the eyelids start to close at the lateral canthus and the closure advances like a zip until it reaches the medial canthus. This flushes tears towards the lacrimal puncta from which point they drain towards the nose.
Upper eyelid margin
Nictitating membrane Many companion pets have a nictitating membrane, or third eyelid, which is a structure located in the medial area and ventral to the eye and held firm by a T-shaped cartilage. The base of the cartilage is connected to the lacrimal gland and both its bulbar and palpebral surfaces are covered with a conjunctiva.The internal surface of the conjunctiva is covered with lymphatic follicles. The nictitating membrane in dogs has a vestigial musculature, which moves passively when the eyeball (innervated by the abducens nerve) is retracted. The lacrimal gland housed in this membrane is important, as it produces 30–50 % of the aqueous component of tears (Fig. 4).
Conjunctiva The conjunctiva is a mobile, elastic mucous membrane covering the internal, or palpebral, surface of the eyelids and the bulbar surface of the nictitating membrane; it is attached to the corneoscleral limbus where it forms the bulbar conjunctiva (Fig. 5). It also has a fornix, which is a cul-de-sac-like structure that establishes the junction between the dorsal and ventral palpebral and bulbar conjunctivae. It is composed of a cylindrical or columnar nonkeratinised epithelium that contains mucus-secreting goblet cells. This mucus corresponds to the protein phase of tears, and amongst other functions, it helps ensure tears adhere to the eye. Any inflammatory or infectious process affecting the ocular surface can alter the conjunctiva’s morphology, causing a loss of goblet cells and modifying its characteristics.
Eyelashes
Lateral canthus Lacrimal caruncle
Free margin of the nictitating membrane
Free margin of the nictitating membrane Conjunctival cul-de-sac (fornix)
Meibomian gland orifices Medial canthus Figure 3. Structures of the eyelid in dogs.
Lower eyelid margin Figure 4. Nictitating membrane and structures of the palpebral conjunctiva.
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