PHYSIOLOGY OF THE EYE

by Khorrami Ph.D.
http://khorrami1962.spaces.live.com khorrami4@yahoo.com http://www.scribd.com/khorrami4

Eye References:

Clinical anatomy of the eye; R.S.Snell Clinical Neuro-ophthalmology; N.R.Miller Adlers physiology of the eye; W.M.Hart Grays anatomy Atlas of strabismus; G.K.V.Noorden Binocular vision & ocular motility; G.K.V.Noorden Ophthalmology; F.W.Newell Neuro-Ophthalmology; Burde Visual perception; S.H. Schwartz

Eye physiology, subjects

Eye movements What is the composition of tear & the mechanisms that control lachrymal glands What are anterior chamber & posterior chamber and which mechanisms can control eye pressure. Iris, pupil & light reflex Composition of cornea & the mechanisms involved in its transparency Composition of lens & the mechanisms involved in its transparency, physiological age related changes of lens Retina & how we can see in light & dark. Retina, cell architecture The pathways where the visual signals pass through the brain How we can sense the colors How we can asses the functions of retina and neural pathways. Optic nerve, afferent & efferent pathways

As sensorimotor system

To see the sights

Sensory system

Vision Depth of vision Color vision Sense of movements

Motor system

Transforming the field of vision into fixation Object onto the fovea Maintenance of binocular vision Connection with vestibular system

Tenons capsule

Muscle characteristics

Diameter Length Mitochondria Reticulum sarcoplasmic, T.tubule Myoglobulin Glycogen Innervation Sensory compartments Fatigability Elastic fibers

Functions of extraocular muscles

Optostatic Optokinetic

Comparison with skeletal muscles

Index EOM Skeletal

Amplitude
Duration of AP Frequency Contraction time

20-150V
1-2ms 150 8ms

100-3000V
5-10ms 50 40-100ms

Innervations
All-or-none

even1-1
yes

1-125
No

Extraocular muscle classification

Kato(1938): based on diameter of fibers

Large
Small

Hess(1963): based on morphology

Fast

twitch( fibrillar type) Slow tonic (field type)

Denervation of EOM (Asmussen 1975)

Atrophy of fast (global) muscles Hypertrophy of slow (orbital) muscles

Extra ocular muscles

PEE & SEE Schiefferdecker:

Abundant

thick parallel elastic fibers Elastic bands

Types of EOM; Peachey(1971)

Connective tissue septa of EOM in orbit

Eye movements

Duction Version Vergence

Eye Movements and Extraocular muscles

Rectus Muscles
Superior and Inferior Oblique

Eye movements

Eye movements

Ocular movements

Dual Actions of Superior Rectus

Dual Actions of Superior Oblique

Action of EOM from primary position

Saccadic eye movements

Fastest movements produced by the human body Quick, simultaneous movements of both eyes in the same direction Frontal and parietal lobes of the brain

Boeders theory(1962)
Vertical movement of horizontal muscles is minimal Horizontal movement of vertical muscles is minimal

EOM; Boeders theory

EOM; Boeders theory

EOM; Boeders theory

EOM; Boeders theory

Eye movements

Pair muscles Yoke muscles

Yoke muscles
Direction Up & right(A) Yoke muscles
RSR + LIO

Right
Down & right(C)

RLR + LMR
RIR + LSO

Down & left(D)

Left Up & left(B) Elevation Depression

RSO + LIR
RMR + LLR RIO + LSR RSR & RIO+LSR & LIO RSO & RIR+LSO & LIR

Lacrimal cells

Light( K cells)
Contain small, electrolucent granules Mucous May function autonomously

Medium

Mixed

Dark( G cells)
Contain large, electron dense granules Serous Cholinergic supply

Lacrimal gland

Tear secretion stimulated by:

Lacrimal nucleus of VII(Ach)

Nicotinic

& muscarinic Emotional Psychic

More

protein-based hormones PRL, ACTH, and leucine enkephalin

Maxillary branch of V(Ach)

Sensory

pathway of reflex
Find & explain

Superior cervical ganglion(sym)

Innervate

mainly blood vessels

Tear layers

Anterior lipid layer(0.2-0.9)

Meibomean,

Zeis & Moll

Middle aqueous layer(6.5-7.5)

Krause,

Wolfring
of cornea

Moistening

Mucinous(0.5 )
Goblet

cells & lacrimal glands

Name

Container(s)

Secretors

Functions coats the aqueous layer; provides a hydrophobic barrier that retards evaporation and prevents tears spilling onto the cheek. These glands are found among the tarsal plates. Thus, the tear fluid deposits and between the eye proper and oil barriers of the lids.[2]

Lipid layer

Oils

Meibomian glands(or tarsal glands)

Aqueous layer

Water and other substances such as proteins (e.g. tear lipocalin, lactoferrin, lysozyme, and llactirin)

Lacrimal gland

promotes spreading of the tear film; promotes the control of infectious agents; promotes osmotic regulation

Mucous layer

Mucin

conjunctival goblet cells

coats the cornea provides a hydrophilic layer; allows for even distribution of the tear film; covers the cornea

Blinking

Shortens canaliculi Expands lacrimal sac Vacuum tears into lacrimal sac Pumping the lacrimal glands Involuntary blinking spreads tear Each 5sec, for 0.3sec Infants have not blinking Infrequent blinking in hyperthyroidism & parkinsonism

Tear inflow & drainage

Drainage of tear

Tears secretion

Normal: 0.9-2.2 l/min Maximum drainage capacity: 30 l/min Crying: 100 l/min Evaporation: 0.85 l/min

Accessory lacrimal glands

Meibomean; sebaceous
10-15

acini

Zeis; sebaceous, connected with follicles Moll; sweat gland Krause; serous, in fornices Wolfring; mucin

Conjunctiva
Areas

Layers

Palpebral
Marginal tarsal Orbital

Stratified columnar epithelium

5-7 layers in corneoscleral 2 layers in tarsal Connective tissue Muscle fibers Blood vessels Nerve, glands Macrophage, mast cells

Sup. & inf. Fornices

Semilunar folds Lacrimal caruncles

Lamina propria

Bulbar

Loosely attached

Corneal layers

Stratified epithelium(10% of tissue thickness)

Consists 1000s tiny nerve endings(very sensitive) Oxidative metabolism Necessary for healing Collagen
Water(78%) + collagen(16%), no blood vessels

Bowman layer

Stroma(Substantia propria), contain keratocytes(90%)

Decemet membrane(single layer)

Collagen, barrier against infection & injuries

Dehydration

Endothelium, microvilli for nourishment

Repair of injury: 6 weeks

Electromagnetic transmit

Wavelengths: 300(UV) 2500nm(IR)

400nm: 80% transmitted 500- 1200nm: 100% Larger than 1000nm cannot stimulate retina

Transparency of the cornea

Anatomic structure Lack of pigments and vessels Cellular arrangements Nucleus and other organelles in periphery Non-keratinized epithelial cells Tight junction, impermeable to aqueous solution Dynamic balance between ions & water Corneal dehydration

Endothelial cells of the cornea

Active transport Passive transport

Epithelial cells of the cornea

35%

65%

Aqueous humor

Maintain IO pressure Nourishment of zonules, lens, iris, cornea & trabecular meshwork Washing the metabolites

Aqueous humor

Production: 2l/min Total volume: 125l By nonpigmented epithelial cells of cilliary body Helps maintain the IO pressure Metabolism of cornea & trabecular meshwork Blood-aqueous barrier: 104 nm
Tight junction in pigment epithelial cells Iris Iris blood vessels

Aqueous humor composition

Na+, K+, Mg2+ as plasma Ca2+ half of plasma Chloride more than plasma Bicarbonate less than plasma Ascorbic acid more than plasma Glutathione more than plasma

Trabecular meshwork

Intra-Ocular pressure

12-20mmHg greater than Atmosphere 2-5mmHg daily fluctuations

Intra-Ocular pressure depends on:

Formation of aqueous humor Trabecular meshwork resistance

75%

of total resistance

Schlemm canal resistance Pressure in episcleral vein(10mmHg)

Blood

pressure Osmotic pressure of blood

Lens layers

Lens; solute movements

Lens metabolism

Glycolysis 85%

Aerobic 5-7% but 25% of ATP

Hexose monophosphate shunt 5-7%

Lens proteins

33% of wet weight of the lens Water soluble, mainly crystallins 80%
Structural Active

signal in lens development

Water insoluble 20%

Cytoskletal
Urea

insoluble

Membrane
Soluble

structural

in 8M urea

Water soluble proteins

Alpha
600-4000kDa
Consist

of 20kDa polypeptides Hydrophobic & H+ bonds

eta 55% of W.soluble

40-200kDa

Gamma
18-20kDa In

embryonic nucleus

Water insoluble proteins

Major intrinsic polypeptide(MIP)

28kDa In gap junctions In differentiating lens fibers Breakdown with aging 28 to 22kDa At ages 20-30 at equal abundance

Glucose on lens
Hyperglycemia

Hypoglycemia

Glycolysis Reduce hexokinase path Shift to sorbitol path Fructose Lens is impermeable to sorbitol and fructose Osmotic pressure Cataract Glycosylation

Reduce ATP Calcium accummulation Protein aggregation Cataract

Transparency of lens(summary)

Is not dependent on oxygen But depends on glucose level in aqueous humor Na+-K+ ATPase activity Fibrillar junctions, sutures Fibrillar proteins Orderly packing of lens fibers Calcium ATPase Na+-Ca2+ exchanger
Finely granular and evenly dense cytoplasm and no organelles

Absence of blood vessels, lymph and nerve Special proteins Dehydration Special metabolism

Retina layers
Bruchs membrane(between RETINA & CHOROID) Retinal pigment epithelium(RPE) Photoreceptors, outer segment Outer limiting membrane( process of Mller cells) Outer nuclear layer(cell body of rod,cone) Outer plexiform layer( ph.R.-neuron synapse) Inner nuclear layer( cell body of Mller) Inner plexiform layer( synapse with ganglion cells) Ganglion cell layer Nerve fiber layer( axon of ganglion) Inner limiting membrane ( process of Mller cells)

Retinal layers

Retina

Macula

Blind spot

Fill-in mechanism

RPE: functions
A part of blood-retinal barrier Takes up nutrients(glucose, retinol & fatty acids) from blood All trans to 11-cis retinal isomerization(visual cycle of retinal) Stabilize ion composition in the subretinal space Phagocytosis of shed photoreceptor outer segments Essential for proper development of the RETINA Survival & differentiation of photoreceptors Secrete a variety of growth factors

Photoreceptors
Rods: apprx. 125mil,
For night vision

Cones: apprx. 6mil,

For central vision

Photoreceptors; distribution

Fovea
1% of retina 110-115K cone cells Yellow; Xanthophyl, Lutein & carotenoids Takes up over 50% of visual cortex Sees only 2o of central visual field(2 thumbnail) Fovea: 1mm, 12/100 m Foveola: 0.2mm, no rod cells, 50/100 m Oxygen supply:
Choroidal vessels More demands under conditions of bright light

Size constancy

Photoreceptor disks
1000 disks in rods Every 30 minutes from base About 3 weeks life Cones for dinner Rods for breakfast

Disc shedding

Photosensors

Photopigments
A protein covalently linked to 11-cis retinal 11-cis retinal is an aldehyde derivative of carotenoid alcohol Opsin in rods contains 348AA, span the membrane 7 times

Rhodopsin

11-cis retinal

Phototransduction
Absorption of a photon by photopigment ( rhodopsin) Prelumirhodopsin Lumirhodopsin Metarhodopsin I-------- Metarhodopsin II/10-3 (active form) Activation of a G-protein(transducin) Activation of phosphodiesterase(PDE) Hydrolyze of cGMP Closing Ca-Na+ channels(300) Hyperpolarization Metarhodopsin II-------- Metarhodopsin III/ several minutes Splits into All-trans retinal and opsin Some converts to all-trans retinol Isomerase from RPE Conversion of all-trans retinal and retinol to 11-cis retinal Attachment of 11-cis retinal to opsin

Photoreceptor

Potential in photoreceptores
Cones 20ms Stimulation rate: 50Hz Rods 50ms Stimulation rate: 20Hz
In cinema: 24frame/sec

Bipolar cell types

Horizontal cells
Ph.R NT depolarize them Hyperpolarize with light C type
Sensitive to color

L type
Sensitive to luminosity or intensity

Inner half of IPL

Outer half of IPL

Cell organization of retina

Facts
Center-surround All photoreceptors are depolarized in dark All photoreceptors hyperpolarized with light Depolarized cells release NT Depolarized bipolar cell stimulate ganglion cell Invaginated BP-cell hyperpolrized with NT of Ph.R. Flat BP-cell depolarized with NT of Ph.R. Depolarized horizontal cell hyperpolarise target Ph.R. Individual Ph.R. synapse with both invag. & flat bipolar Invaginating bipolar just receive one type of synapse

Ganglion cell

Children of the light

Color opponent ganglion cells

Ganglion cells
Types X (P) Y(M) 80%, sustained response 10%, transient response 10%, transient response Importance Visual acuity, color, fine details Shape of objects, contrast, low resolutionrsg Luminosity, head & neck movements Receptive field Smallrsg Largesg Projection LGN LGN, sup. coll

Largesg

Sup. coll

Ganglion cells

Perimetry

Retinofugal projections
Retinogeniculostriate pathway Extra retinogeniculostriate pathway
Retinotectal(superior colliculus) to:
R.F. in arousal response LGN . . . Cortex

Pretectal Pregeniculate nucleuspontine nucleusmossy fiber.vestibulocerebellum Accessory optic nucleus.inferior olive N.climbing fibervestibulocerebellum SCNcircadian rhythm/ melanopsin in a type of ganglion cells

Retinal projections

LGN & projections to cortex

Central processing
Orientation selective neurons, by edge detecting What pathway: parvocellular, fine, color and dept
From P-type cells Slowly adapt V1,V2,V4 & IT IT(inferotemporal) cortex; visual long term memory
Stimulated in epilepsy

Where pathway: magnocellular: motion, position

Rapidly adapt V1,V2,V3, V5(medial temporal) Finally to medial temporal, medial superior temporal & posterior parietal(Brodman5&7 )

Dorsal and ventral stream

IT: inferotemporal PP: posterior parietal ST: superior temporal

Striate cortex
M channel
Analysis of object motion

P-interblob channel
Analysis of object shape

Blob channel
Analysis of object color If defect . . . . Color blindness

Layer VI . . . . To LGN for feedback Layer V . . . . To superior colliculus for visual reflex Layer II & III . . . . Between cortex

Postretinal pathways
Magno retinogeniculate pathway:
Motion

Parvo retinogeniculate pathway:

Red-green color vision

Konio retinogeniculate pathway:

Blue-Yellow color vision

Eye movements
With head movements
Vestibulo-ocular eye movements Opto-kinetic eye movements

No movement in head
Saccadic eye movements (vertical or horizontal) Smooth pursuit eye movements Vergence eye movements

Eye movements
Superior muscles receive innervations from opposite side PPRP in horizontal movements
Receive from: sup.col., Vest.N., cerebellum, frontal eye field, MLF, perihypoglossal N.( in gaze holding) Ipsilateral LR & contralateral MR Left side lesion.cannot follow the object in left

riMLF in vertical movements Vermis of cerebellum in generation of REM Superior colliculus

Color perception
Rhodopsin on chromosome 3 Blue(tritan) on chromosome 7 Red(protan) and green(deutan) on chromosome X Protanomaly: poor red-green, dimmer red Deutranomaly: poor red green, normal red Protanopsia Deutranopsia

Normal color vision

Protanopia

Deuteranopia

Tritanopia

Forebrain Control of Eye Movements

Multiple paths descend from forebrain and affect eye movements. These paths project either directly to brainstem nuclei involved with eye movements or relay via the superior colliculus. Frontal eye fields appear to be in area 6. FEF generates saccades in the contralateral direction via connections to contralateral PPRF. Parieto-occipital-temporal cortex functions in smooth pursuit eye movements in ipsilateral direction via connections with the vestibular nuclei, cerebellum, and PPRF. Inputs from visual cortex and visual association cortex influence FEF activity. Basal ganglia also appear to play a role in eye movements.

Visual location

Motion( middle temporal gyrus)

Achromatopsia( occipito-temporal gyrus)

Visual agnosia( posterior & medial temporo-occipital cortex)

Visual neglect( inferior parietal lobe)

Balint-Holmes syndrome*( posterior parietal lobe)

Optic ataxia, ocular apraxia & simultanagnosia

Visual cortices
A: visual location B:Motion blindness(middle temporal) C: Achromatopsia (occipito-temporal) D: Visual agnosia ( post. & medial temporo-occipital) E: Visual neglect (inf. parietal) F: Balint-Holmes syd.(post. Parietal)