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Ординатура / Офтальмология / Английские материалы / Imaging of Orbital and Visual Pathway Pathology_Muller-Forell_2005

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52

W. Wichmann and W. Müller-Forell

Fig. 2.37. Axial T1-weighted MRI of the optic nerve, demonstrating the tortuous course of the left as a normal variant

2.7

Intracranial Visual Pathway

2.7.1

Optic Tract (Figs. 2.38, 2.39)

Dorsolaterally of the chiasm, the optic visual pathway continues backwards, forming the optic tracts. They cross posterior to the anterior perforated substance and anterior to the lateral tuber cinereum and the lateral perforated substance, respectively (Duvernoy 1998). More dorsolaterally, the optic tracts lie above the uncus and surround the crus cerebri. The main part of the tracts ends in the lateral geniculate nuclei (Figs. 2.43, 2.44). Small divisions of the tracts terminate in the superior colliculus, in the pretectal area, in nuclei of the accessory optic tract, and in the hypothalamic suprachiasmatic nucleus (Nieuwenhuys et al. 1988). The brachium colliculi superiores, which courses below the lateral pulvinar of the thalamus between the lateral geniculate nucleus and the colliculus superior, also contains others fibers that connect the lateral geniculate nucleus with the colliculus superior. On high resolution MRI, the lateral geniculate nucleus and the brachium colliculi superioris (Fig. 2.44) can be depicted (Nieuwenhuys et al. 1988; Horton et al. 1990; Duvernoy 1998).

(Text continues on p. 55)

12.1

14.9

14.9

12.2

ant. choroid. artery

12.3

12.3

13.21

13.13

13.15

15.2

12.6

15.3

a

b

Fig. 2.38.a Axial T2-weighted MRI of the chiasmatic region. b Corresponding diagram: 12.1 = prechiasmatic optic nerve, 12.2 = chiasm, 12.3 = optic tract, 12.6 = optic radiation, 13.13 = mammillary body, 13.15 = pineal gland, 13.21 = third ventricle, 14.9 = MCA (M1), 15.2 = (distal portion of the ) internal cerebral veins (draining into the), 15.3 = (proximal) vein of Galen (note the wide prechiasmatic cistern (as an individual, normal variant), especially in comparison with Fig. 2.39)

Anatomy

53

ant. comm. artery (double)

14.9

 

 

 

14.8

14.8

 

 

 

 

14.9

12.3

13.21

12.3

 

 

 

14.6

 

 

14.7

 

 

13.6

a

 

b

Fig. 2.39.a Axial T2-weighted (MIN) view of the chiasmatic region. b Corresponding diagram: 12.3 = optic tract, 13.6 = pons, 13.21 = third ventricle, 14.6 = posterior communicating artery (Pcomm), 14.7 = PCA (P1), 14.8 = ACA (A1, A2), 14.9 = MCA (M1)

 

14.8

 

14.4

 

14.4

14.2

 

14.2

 

 

14.9

14.6

14.9

 

 

 

ant. comm. artery

 

 

14.1

 

a

 

b

Fig. 2.40.a Axial view of 3D-MRA of the anterior part of the circle of Willis. b Corresponding diagram: 14.1 = (ICA in the) carotid canal, 14.2 = ICA, 14.4 = ophthalmic artery, 14.6 = Pcomm, 14.8 = ACA (A1, A2), 14.9 = MCA (M1–M3)

54

 

W. Wichmann and W. Müller-Forell

14.4

14.8

 

14.4

 

 

 

 

14.3

 

 

14.3

 

14.9

14.6

 

 

14.5

 

 

 

 

14.9

 

 

 

 

 

 

 

 

 

 

 

14.7

 

 

 

 

15.2

 

 

a

 

 

 

 

b

Fig. 2.41.a 2D-MRA of the circle of Willis. b Corresponding diagram: 14.3 = siphon of ICA, 14.4 = ophthalmic artery, 14.5 = (top of the) basilar artery, 14.6 = Pcomm, 14.7 = PCA, 14.8 = ACA (A1, A2), 14.9 = MCA (M1–M3), 15.2 = (distal portion of the) internal cerebral vein

13.8

13.17

13.14

15.2

 

 

 

 

 

 

15.3

 

13.20

13.10

 

ant. commissure

 

13.15

 

 

 

 

12.2

13.21

 

 

 

 

 

13.12

 

13.26

 

 

 

 

 

13.13

 

3.12

13.11

13.6

 

 

 

 

 

 

13.25

a b

Fig. 2.42.a Midsagittal T1-weighted MRI of the sellar region. b Corresponding diagram: 3.12 = sphenoid sinus, 12.2 = chiasm, 13.6 = pons, 13.8 = rostrum (of the corpus callosum), 13.10 = thalamus, 13.11 = pituitary gland [anterior pituitary (adenohypophysis): isointense, posterior pituitary (neurohypophysis): hyperintense], 13.12 = pituitary stalk, 13.13 = mammillary body, 13.14 = fornix, 13.15 = pineal gland, 13.17 = ventricle, 13.20 = foramen of Monro, 13.21 = third ventricle, 13.25 = fourth ventricle, 13.26 = aqueduct, 15.2 = internal cerebral vein, 15.3 = vein of Galen

12.7

 

 

substantia nigra

 

12.5

 

 

12.4

12.5

 

 

 

 

13.26

a

 

b

Fig. 2.43.a Axial PD-weighted MRI of the mesencephalic region (sector) at the level of the anterior commissure. b Corresponding diagram: 12.4 = medial geniculate nucleus, 12.5 = lateral geniculate nucleus, 12.7 = anterior commissure, 13.26 = aqueduct

Anatomy

 

 

55

13.17

13.9

13.17

 

13.14

 

13.14

 

13.10

 

13.10

12.4

 

 

12.5

brachium colliculi sup

13.19

 

 

13.19

 

 

13.16

 

 

 

tentorium

 

a

 

13.25

b

Fig. 2.44.a Coronal T1-weighted (IR) view of the midthalamic region. b Corresponding diagram: 12.4 = medial geniculate nucleus, 12.5 = lateral geniculate nucleus13.9 = body (of the corpus callosum), 13.10 = thalamus, 13.14 = fornix, 13.16 = quadrigeminal plate, 13.17 = ventricle, 13.19 = temporal horn of the ventricle, 13.25 = fourth ventricle

2.7.2

Optic Radiation (Figs. 2.45, 2.46)

About half of the fibers that originate in the lateral geniculate nucleus and project to the occipital visual cortex run directly backward lateral to the more superior aspect of the lateral ventricle within the inferior parietal lobe. The other half of the fibers describes a large forward directed loop (Meyer’s loop) within the temporal lobe above and lateral to the anterior temporal horn of the lateral ventricle. Then the fibers curve backward lateral to the temporal and occipital horn of the lateral ventricle. The fibers of the optic radiation are separated by the interposed tapetum from the lateral wall of the ventricle. The tapetum is a thin layer composed of fibers from the forceps major of the callosal radiation. Tapetum and optic radiation together form a thin caudo-cranially directed plate of fibers, termed the sagittal stratum. On T2-weighted MRI, it is possible to identify and trace systems of parallel coursing, compact, myelinated fibers as the optic radiation, which are embedded in more loosely arranged white matter (Curnes et al. 1988). On MRI in most cases, it is not possible to subdivide the sagittal stratum and differentiate the optic radiation and the tapetum. Only under special conditions, if there is edema or a gliosis, can they be separated on T2-weighted MRI (Kitajima et al. 1996). Behind the sagittal stratum, the myelinated fibers of the optic radiation bend medially to project on the visual cortical field and diverge in a fan-like manner, no longer depicted on MRI as the fibers are looser than the optic radiation.

Fig. 2.45.a Axial T2-weighted view of the brain at the level of the internal capsule. b Corresponding diagram: 12.6 = optic radiation, 13.10 = thalamus, 13.18 = frontal horns of the ventricles, 13.20 = foramen of Monro, 13.21 = third ventricle, 15.3 = vein of Galen

a

13.18

13.18

internal capsule

internal capsule

13.20 13.20

13.21

13.1013.10

 

15.3

12.6

12.6

b

56

W. Wichmann and W. Müller-Forell

a

13.14

 

 

 

13.14

 

 

13.7

 

 

 

 

 

 

 

 

12.6

 

 

 

 

12.6

 

 

 

13.313.3

b

Fig. 2.46.a Axial PD-weighted view of the optic radiation some millimeters above Fig. 2.45. b Corresponding diagram: 12.6 = optic radiation, 13.3 = occipital lobe, 13.7 = (posterior knee of the ) corpus callosum, 13.14 = fornix

contralateral visual field shows its representation below the horizontal meridian. The peripheral visual field is projected to the anterior part of this ellipsoid map where the calcarine fissure joins the parietooccipital fissure. The striate cortex around and lateral to the occipital pole contains the representation of the central visual field, especially of the fovea. An expanded area of the visual cortex is attributed to the central visual field, and a relatively small part is occupied by the peripheral visual field representation. This anatomic arrangement is called cortical magnification of the central vision (Holmes 1945). Correlative studies of the past few years, since functional MRI has been able to localize occipital lesions with attributed visual field defects in an anatomical way, have led to a postulated refinement of the classic retinotopic Holmes map. It seems that the abovementioned so-called cortical magnification of the central visual field was underestimated by Holmes (Horton and Hoyt 1991b; Wong and Shape 1999). A revised map of the retinotopic representation of the human visual striate cortex is shown in Figs. 2.49, 2.50 (Horton and Hoyt 1991a). According to the literature, high resolution MRI is able to identify in vivo the specific intracortical myeloarchitecture of the striate cortex that differs from the extrastriate cortex (Clark et al. 1992). This can be demonstrated by comparison of the striate and precentral cortex obtained from specimens (Fig. 2.51), where the myeloarchitecture shows a more distinct visibility than on in vivo MRI.

2.7.4

Extrastriate Visual Association Cortex

2.7.3

Striate Cortex (Figs. 2.47–2.50)

The primary visual cortex, the striate cortex, is located in the wall along the calcarine fissures, seen on the medial surface of the occipital lobe (Figs. 2.47, 2.48). It should be emphasized that the striate cortex wraps around the occipital pole to the posterolateral aspect of the occipital lobe. In other words, one can imagine folding back the calcarine fissure laterally and smoothing out the normally gyrated striate cortex, leading to an artificially flattened ellipsoid map of the visual cortex that is about 80–40 mm (Horton and Hoyt 1991b). The folding along the calcarine fissure indicates the horizontal meridian. The lower contralateral visual field is represented above the horizontal meridian, whereas the upper

Except anteriorly, the primary visual cortex (V1 or area 17) is surrounded by the secondary visual area V2 (area 18), which in turn is surrounded by the tertiary visual area V3 (area 19 corresponds partially to V3). These areas are designated the circumstriatal visual association cortex. Whereas the upper and lower quadrants of the primary visual cortex are in continuation within the depth of the calcarine fissure, the extrastriate cortex is divided into separate upper and lower quadrants flanking above and below the intervening primary visual cortex. Each quadrant of the bisected secondary visual cortex runs on both sides of the primary visual cortex along the upper and lower vertical meridians which are shared by the primary and secondary cortex. On the outer border of the secondary cortex, the quadrants of the bisected tertiary cortex are arranged along the horizontal

Anatomy

57

parieto-occipital sulcus

13.5

13.5

13.4 13.4

13.3

13.17

13.3

tentorium

a

b

Fig. 2.47.a Coronal T1-weighted view (reconstruction of a 3D data set) of the region of the calcarine sulcus. b Corresponding diagram: 13.3 = occipital lobe, 13.4 = calcarine sulcus, 13.5 = parietal lobe, 13.17 = (most posterior parts of the inferior horns of the) ventricles

parieto-occipital sulcus

15.2

13.7

13.4

 

 

 

 

15.3

 

 

13.16

tentorium

 

 

13.6

13.25

c

d

Fig. 2.48.a Sagittal paramedian T1-weighted (IR) view of the parieto-occipital region with striate cortex. b Corresponding diagram: 13.4 = calcarine sulcus, 13.6 = pons, 13.7 = corpus callosum, 13.16 = quadrigeminal plate, 13.25 = fourth ventricle, 15.2 = internal cerebral vein, 15.3 = vein of Galen

meridian that is shared by the adjoining secondary and tertiary cortex (Fig. 2.52). This explains why a lesion of the extrastriate cortex that crosses the horizontal meridian between areas V2 and V3 will respect the horizontal meridian and result in a perfect quadrantanopia. In this described condition, a precise

anatomical location or extension of the lesion is not required. In the case of a quadrantanopia, this does not necessarily imply that a lesion lies within the extrastriate cortex; it can also be caused indirectly by compression by a bordering tumor (Van Essen et al. 1986; Horton and Hoyt 1991a).

58

W. Wichmann and W. Müller-Forell

Fig. 2.49. Schematic diagram showing arrangement of V1, V2, and V3 along the medial and posterior occipital surface. Most of V1 is buried within the calcarine fissure. Considerable variation occurs among individuals in the relative position and size of different cortical visual areas. (With permission of Horton and Hoyt 1991a)

 

 

 

 

a

 

 

 

 

b

Fig. 2.50.a Artificially flattened map showing the retinotopic organization of V1 (stippled area), V2 (small triangles), and V3 (striped) in the left occipital lobe. b Right visual field coordinates corresponding to map in a. The monocular temporal crescent (stippled area) is represented within a small area at the rostral end of striate cortex (border between binocular and monocular field runs between arrows in a). V2 and V3 are split along the representation of the horizontal meridian into separate dorsal and ventral halves. More than half of the visual cortex is devoted to processing the central 10º of vision. (With permission of

Horton and Hoyt 1991a)

 

Gennari-stripe

 

white substance

 

inner cortex

 

with myelin-fibers

 

striate cortex

a

sulcus

b

Fig. 2.51.a PD-weighted high resolution MRI (3.8 T) of a human specimen (1.5 mm thickness) of the striate cortex. b Corresponding diagram (the compartments are shown lateral to the white substance)

Anatomy

Fig. 2.52. Schematic coronal section through right occipital lobe showing connections from V1 (stippled area) to V2/V3 (small triangles and striped, respectively). Projections pass between common retinotopic coordinates. In V1, the horizontal meridian is represented along the base of the calcarine sulcus. Fibers coursing in the white matter of the upper and lower calcarine banks serve the lower and upper visual quadrants, respectively. For simplicity, feedback pathways from V2/V3 to V1 and projections between V2 and V3 are omitted. The ventral V1 and V3 pathway is doubtful (Van Essen et al. 1986). (With permission of Horton and Hoyt 1991a)

References

Breslau J, Dalley RW, Tsuruda JS, Hayes CE, Maravilla KR (1995) Phased-array surface coil MR of the orbits and optic nerves. Am J Neuroradiol 16:1247–1251

Bron AJ, Tripathi RC, Tripathi BJ (1997) Wolff’s anatomy of the eye and orbit, 8th edn. Chapman and Hall, London

Casper DS, Chi TL, Trokel SL (1993) Orbital disease. Imaging and analysis. Thieme, New York

Clark VP, Courchesne E, Grafe M (1992) In vivo myeloarchitectonic analysis of human striate and extrastriate cortex using magnetic resonance imaging. Cereb Cortex 2:417–424

Curnes JT, Burger PC, Djang WT, Boyko OB (1988) MR imag-

59

ing of compact white matter pathways. AJNR Am J Neuroradiol 9:1061–1068

Daniels DL, Leighton PM, Mafee MF et al (1995) Anatomic moment. osseous anatomy of the orbital apex. AJNR Am J Neuroradiol 16:1929–1935

Duvernoy HM (1998) The human hippocampus, 2nd edn. Springer, Berlin Heidelberg New York

Helmke K, Hansen HC (1996) Fundamentals of transorbital sonographic evaluation of optic nerve sheath expansion under intracranial hypertension. I. Experimental study. Pediatr Radiol 26:701–705

Hoffmann KT, Hosten N, Lemke AJ, Sander B, Zwicker C, Felix R (1998) Septum orbitale: high-resolution MR in orbital anatomy. AJNR Am J Neuroradiol 19:91–94

Holmes G (1945) The organization of the visual cortex in man. Proc R Soc Lond B 132:348–361

Horton JC, Hoyt WF (1991a) Quadrantic visual field defects. Brain 114:1703–1718

Horton JC, Hoyt WF (1991b) The representation of the visual field in human striate cortex. A revision of the classical Holmes map. Arch Ophthalmol 109:816–824

Horton JC, Landau K, Maeder P, Hoyt WF (1990) Magnetic resonance imaging of the human lateral geniculate body. Arch Neurol 47:1201–1206

Kitajima M, Korogi Y, Takahashi M, Eto K (1996) MR signal intensity of the optic radiation. AJNR Am J Neuroradiol 17:1379–1383

Lang J (1981) Klinische Anatomie des Kopfes. Springer, Berlin Heidelberg New York, p 92

Nieuwenhuys R, Voogd J, van Huijzen C (1988) The human central nervous system, 3rd edn. Springer, Berlin Heidelberg New York, pp 20, 92, 107, 182, 183

Oikawa S, Kyoshima K, Kobayashi S (1998) Surgical anatomy of the juxta-dural ring area. J Neurosurg 89:250–254

Renn WH, Rhoton AL Jr (1975) Microsurgical anatomy of the sellar region. J Neurosurg 43:288–298

Rhoton AL, Natori Y (1996) The orbit and sellar region: microsurgical anatomy and operative approaches. Thieme, New York

Servo A (1982) Visualization of the superior ophthalmic vein on carotid angiography. Neuroradiology 23:141–146

Van Essen DC, Newsome WT, Maunsell JHR, Bixby JL (1986) The projections from striate cortex (V1) to areas V2 and V3 in the macaque monkey: asymmetries, areal boundaries, and patchy connections. J Comp Neurol 244:451–480

Williams PL (1995) Gray’s anatomy, 38th edn. Churchill Livingstone, New York

Wong AM, Shape JA (1999) Representation of the visual field in the human occipital cortex: a magnetic imaging and perimetric correlation. Arch Ophthalmol 117:208–217

Neuro-ophthalmology: A Short Primer

61

3Neuro-ophthalmology: A Short Primer

Urs Schwarz

CONTENTS

3.1Introduction 61

3.2

Vision and Visual Perception 61

3.2.1Receptive Fields 64

3.2.2

The Retino-geniculo-cortical Pathway 67

3.2.2.1

The Retina 67

 

 

 

3.2.2.2

The Lateral Geniculate Nucleus

67

3.2.2.3

The Primary Visual Cortex

68

 

3.2.2.4

Lesions in the Retino-geniculo-cortical Pathway 70

3.2.3

The Extrastriate Visual Cortex

72

3.2.3.1

The Dorsal Pathway

72

 

 

3.2.3.2

The Ventral Pathway

73

 

 

3.2.4

The Accessory Optic System

74

3.3Eye Movements 74

3.3.1The Oculomotor Nuclei and the Extraocular Muscles 74

3.3.1.1

Anatomy of the Ocular Motor Nuclei and Nerves

76

3.3.1.2

Lesions of the Extraocular Nuclei and Nerves

78

3.3.2Supranuclear Synchronization of Eye Movements 84

3.3.3

Gaze Holding Mechanisms

87

3.3.3.1

The Vestibulo-ocular Reflex

87

3.3.3.2

The Optokinetic Reflex 92

 

3.3.3.3

Nystagmus

92

 

3.3.4

Gaze Shifting Mechanisms

93

3.3.4.1

Saccades

93

 

3.3.4.2

Smooth Pursuit Eye Movements 97

3.3.4.3Vergence 98

3.3.4.4Fixation 98 References 98

3.1 Introduction

The spectacular symbiosis between the visual system, the vestibular system, and the optomotor system, each of which is a miracle on its own, is able to solve an abundant array of amazing tasks (Fig. 3.1). In particular, the connection between vision and the optomotor control is most intriguing, as the eye performsboththeinputandtheoutputsimultaneously. Altogether,it is not surprising that the implementation of these intricate neural networks encompasses almost

every part of the brain (Fig. 3.2). Moreover, these systems are probably amongst the best investigated so far; and enormous knowledge has been amassed over the past few decades employing a variety of techniques from single cell recordings, optical imaging, eye movement recordings, functional magnetic resonance imaging, to neuropsychological examinations. In addition,this research has clarified many fundamental principles of neural and neuronal processing, which subsequently have enriched other fields of brain research as well as computational neuroscience.

The optomotor system provides a magnificent windowintothenervoussystemforclinicians,aspeople suffering from even the smallest disruption soon are unable to pursue their regular routines. Armed with thorough neuroanatomical and neurophysiological knowledge of this system’s workings,a clinician will be able to diagnose and localize many conditions right at the bedside.

A systematic review bearing on the whole spectrum of normal and pathological visual,vestibular,and optomotor neural processing would be more than one’s lifetime work. Hence, this improperly short primer is very selective, fractional, and undeniably biased. An attempt was made to seduce the reader to dive further into this fascinating realm of neuro-ophthalmology by presenting a basic framework of functional neuroanatomy. The basic neuroscientist will find plenty in the works of Carpenter (1988), Zeki (1993), Milner and Goodale (1995), Wandell (1995), Zigmond et al. (1999), Kandel et al. (2000); the clinician, on the other hand, will find more answers in the seminal oeuvres (Baloh and Honrubia 1990; Grüsser and Landis 1991; Miller and Newman 1997; Huber and

Kömpf 1998; Leigh and Zee 1999).

3.2

Vision and Visual Perception

U. Schwarz, MD

Light entering the eye is projected onto the retina,

Priv.-Doz., Department of Neurology, University Hospital

(USZ), Frauenklinikstrasse 26, 8091 Zürich, Switzerland

where it is converted into an electrical signal by the

62 U. Schwarz

vestibulartasks

visualtasks

 

 

 

unexpected

 

 

 

 

 

 

 

 

disruptionof

 

 

 

 

posture

 

 

 

 

 

 

 

 

 

 

moving

target

vL

 

 

visual

-selection

 

suroundings

 

internal

 

-recognition

 

F

 

representation

 

maintenanceof

 

selected

 

egomotion

gaze

spatialorientation

 

target

between

 

 

-self

 

 

-target

 

next

 

 

possible

-visualsuroundings

 

 

 

 

target

Fig. 3.1. Optomotor tasks. The visuo-, vestibulo-, and proprioceptive-optomotor systems ensure maintenance of orientation in space as well as target selection and recognition despite a host of sensory conflicts and adversary disturbances. Its main goals are to keep the target of interest on the fovea (stabilization of gaze) and to produce an accurate internal representation of spatial relations between the visual surroundings, the target, and the self. vL, vestibular labyrinth

 

aference

supranuclear

 

eference

 

 

control

 

 

SP

 

 

 

 

 

 

epS

sC

aoS

VI VI

 

 

 

 

IV

IV

 

 

 

 

 

vT

 

 

PPRF

riMLF

 

 

 

 

 

 

I I

 

vis

 

 

 

conjugate

 

 

 

MLF

 

 

OKR

Cbl

 

 

 

and

 

 

 

 

 

vest

 

 

 

vergence

 

pvS

 

 

 

 

 

 

 

 

 

vS

 

NV

NV

 

 

 

 

gazepalsy

gazepalsy

 

diplopia

inappropriateeyemovements INO

disintegratedeyemovements

saccadedeficits

Fig. 3.2. Synopsis of the optomotor system (modified after Schwarz et al. 2000). Multisensory, visual, vestibular, and proprioceptive (not shown) signals (afference) are cortically and subcortically integrated and guided to the supranuclear optomotor control system, which computes appropriate conjugate and vergence signals for each eye muscle and distributes them via the medial longitudinal fasciculus (MLF) to the individual oculomotor nuclei (efference). Therefore, the supranuclear control implements the final common pathway for any mechanism that requests eye movements. Typical eye movement patterns due to lesions of the afferences, supranuclear control, and efferences are shown at the bottom. The exemplary scene shows the complexity of the visual computation: While smoothly following a moving visual target (vT) with the eyes, the visual surroundings (vS) is shifted in the opposite direction (evoking full field stimulation) and, hence, elicits an optokinetic response (OKR). Sensory integration must solve the problem of these counteracting demands and generate appropriate signals to the supranuclear control for smooth pursuit eye movements (SP) only. pvS, peripheral vestibular system; epS, extrapyramidal system; aoS, accessory optic system; Cbl, cerebellum; NV, vestibular nucleus; PPRF, paramedian pontine reticular formation; riMLF, rostral interstitial nucleus of the MLF; III, oculomotor nucleus; IV, trochlear nucleus; VI, abducens nucleus; INO, internuclear ophthalmoplegia

photoreceptors.A geometrically well-defined array of these highly specialized cells connects via an intrinsic local network to one single ganglion cell, thus implementing the first receptive field of the visual system. Consecutively, signals of the retinal ganglion cells are transmitted through their axons, which together form the optic nerve, to the optic chiasm in front of the pituitary stalk, where half of them are

cross routed: Axons from the temporal half of one retina join axons from the nasal part of the other retina in the optic tract. Axons from the temporal retina remain ipsilateral. Hence, visual objects in one hemifield, which are projected nasally onto the ipsilateral retina and temporally onto the contralateral retina, are processed in contralateral central visual centers with information coming from both retinae.

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