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69 The Pupillary Reaction to Near Vision As in the sections on the reactions to light, the pupillary reaction to near vision is considered in its physiological aspects only, and work dealing with the optical effects of pupil size during near vision has been excluded from this review. In Table 9, the publications available to the author have been listed. In contrast to the many aspects of light reactions, not much can be said about this literature. The greater part of the discussions have revolved around the question of whether the pupillary contraction depends on con- vergence or on accommodation. These discussions are still current, even though it has been established that any one of the three functions can be abolished without interfering with the others. Thus, convergence or accommodation may be eliminated selectively by the use of prisms or lenses. Clinical cases with isolated impairment or loss of pupillary contraction, accommodation, or convergence are not too uncommon, and in experiments on animals, isolated pupillary constriction, accommoda- tion of the lens, and contraction of the internal rectus muscle were ob- â¢tained by electrical stimulation in the cortex, the oculomotor nuclear complex, or the efferent third nerve. These facts lead to the conclusion that the nervous impulses that cause accommodation, convergence, and pupillary constriction must arise from different cell groups within the third nerve nucleus, and travel via separate fibers to their effector muscles, whereby the paths for the ciliary muscle and the iris sphincter undergo synapses in the ciliary ganglion. The theory has been proposed that the pupillary fibers for near vision do not synapse in this ganglion while those for the light reflex do, thus explaining the isolated loss of the pupillary light reflex in the pres- ence of unimpaired pupillary contraction to near vision in the Argyll Robertson syndrome (Naquin, 1954). This theory is disproven by the fact that in monkeys, retrobulbar injection of a ganglion-blocking agent such as nicotine entirely abolishes the pupillary contractions elicited by intracranial stimulation of the third nerve with strong currents (Loewenfeld, 1958). Accommodation, convergence, and pupillary contraction are asso- ciated movements, and are not tied to one another in the manner usually referred to by the term "reflex". They are controlled, synchronized, and associated by supranuclear connections, and are not caused by one another. As a final point, it must be remembered that the pupillary reaction to near vision is subject to the same central nervous inhibitory and sympathetic antagonistic influences as are the contractions to light. The extent of the reactions may, therefore, vary from one moment to the other. Just as for the light reflex, these variations have given rise to some unusual statements in the literature. For example, von Graefe's
70 TABLE 9 Pupillary Movements and Near Vision E â¢"' Fusion S 5 irn -in Other Â« on i E H Clinical Animals Year Author Â£ Findings and Conclusions 1619 Scheiner * first to describe pupillary contraction to near vision 1659 Plempius * similar description as Scheiner's 1780 Olbers Â« measured pupil contraction with accommodation method of Lambert 1821,51 Weber * * near vision contracts pupil even if object is darker; accommodation alone does not cause miosis, con- vergence does 1835 Plateau * learned to accommodate without converging: pupil contracted 1853 Cramer * accommodative tension without convergence may contract pupil 1853 De Ruiter * accommodation alone may contract pupil 1854 von Graefe * reported observation of Muller: in dog, pupil dilates upon near vision 1856 von Graefe * case with complete extraocular paralysis; vision perfect; pupil immobile to light but contracted well to near vision; concl. : pupil reaction does not de- pend on extraocular movements; probably mid- brain lesion 1864, 65 Donders * * pupil may contract with accommodation alone or with convergence alone 1866 Trautvetter * repeated test on dog and cat; same result for dog (dilation); cat uncooperative 1868 He ring * entoptic images fused without convergence*-pupil contraction; concl. : accommodation alone may cause contraction, but the three movements are asso ciated, not interdependent 1869 von Arlt â¢ pupil contraction occurs later than accommodation 1869 Le Conte * learned to accommodate without converging*-pupil constricted; concl.: pupil is associated with accom- modation but may also constrict with strong con- vergence alone 1870 Adamiik * obtained associated pupil contraction, movement of globes in & down, from anterior midbrain of dog; pupil may contract with accommodation in absence of convergence 1871 Adamuk t Woinow * measured depth of anterior chamber, lens curva- ture (dog); electrical stimulus to efferent nerves 1873 Coccius * accommodation alone and convergence alone may contract pupil 1874 Kreuchel (i Mulder) 1 studied effect of muscarine on accommodation and pupil 1876 Drouin 1 pupil and accommodation are associated but not dependent on one another
71 TABLE 9 (Cont'd) Lenses PI Prisms Â§ 3 rir it-n BUI ta ii M c bO Clinical Animals 0 1 H i Year Author 6 Findings and Conclusions 1878 H i> n sen 4 V Bickers * obtained isolated accommodation, pupil constriction, contr. medial rectus, from floor of Sylvian atjueduct (dog); after death, extraocular reactions to brain stimuli elicited 15 min. longer than pupil and accommodation 1880 Angelucci & Aubert * measured speed of pupil and of movement of Purkinje images: pupil was slower 1888 Donders (2nd ed.) * * neither iris nor accommodation are dependent on convergence 1892 Borthen * * * case with absent light reflex, good accommodation, pupil contraction only with convergence; concl.: pupil is relatively independent of accommodation 1893 Du Bois-Reymond & Greeff * near vision effort in total darkness (flash photograph): pupil contracted 1898 Heine * * in birds, pupil contraction faster than accommodation (electrical stimulus efferent nerves) 1898 Hess & Heine * dog, cat, rabbit, even when young, have poor accommodation 1898 Ovio tested pupil, accommodation, convergence with one or both eyes open; concl. : pupil associated with both accommodation and convergence, depends on distance of vision 1900 Vervoort * of convergence 1902. 03 Marina * exchanged internal and external recti in monkeys: pupil contraction with convergence post-operative = with contraction of external recti; concl.: no rigid connection in nervous centers 1903 Friberger * measured speed of pupillary movements associated with convergence and accommodation 1904 Bach * cases with (a) complete loss of accommodation, (b) paralysis of internal rectus, (c) unilateral amblyopia or loss of eye: pupil near vision reaction normal; concl.: accommodation, pupil, convergence are asso- ciated movements but not dependent on one another 1905 Moderow * analysis of literature; concl.: convergence is, but accommodation is not associated with pupil 1905 Ovio not read by reviewer 1963 1905 Wlotzka * accommodation alone: pupil immobile; concl. : pupil, accommodation both depend on convergence 1908 Kanngiesser â¢ * pupil contraction with accommodation alone; atropine affected accommodation longer than pupil 1908 Lohmann * accommodation changed without convergence; case with post-diphtheric loss of accommodation; concl. : pupil contraction is synergic co-movement with accommodation and convergence 1909 Magitot * pupil contraction works in 5th foetal month, accommo- dation only months after birth
72 TABLE 9 (Cont'd) E Prisms 3 "g nil M en â¢ in n I 1 B 4 | TJ Clinical Animals B h I i Year Author e E c Findings and Conclusions 1912 Hesse * cases with extraocular paralysis could not converge: pupil contraction with accommodation 1912 IsakowiU * critique of Hesse: absence of convergence movement does not prove absence of convergence impulse 1920 Pick * 8 diopters accommodation without convergence: pupil immobile; pupil is synergic co-movement 1920 Sattler * in high, uncorrected myopes pupils contract when eyes converge without accommodation 1922 Caspary & Goeritz * obtained changes in accommodation without pupil contraction; concl.: pupil tied to convergence, not to accommodation; normally, the three functions occur together 1926 Holladay * pupils contract to forced convergence = chief cause of near vision contraction 1928-29 Kestenbaum & Eidelberg * * * diverse cases, quoted second-hand; concl.: accom- modation and pupil work together only in presence of convergence; pupil = associated movement (with convergence) 1930,31 Gualdi I 1 accommodation alone does not affect pupil, conver- gence alone does 1936 Schubert & Burian * find reflex to fusion, independent of convergence or accommodation 1937 Haessler * tested near vision in near-darkness (luminous dial): pupil contracted 1943 Bender & Weinstein * obtained isolated contraction internal rectus, miosis, accommodation by electrical stimulation in third nerve nuclear complex (monkey) 1945 Fry * complex apparatus, allowing independent control of convergence and accommodation; concl. : pupil, ac- commodative vergence and accommodation are asso- ciated movements; influence of fusional convergence questionable 1949 Knoll * complex apparatus; pupil constricted with accommo- dation and accommodative vergence; in some subjects pupil also constricted with fusional convergence 1949 Marg & Morgan 1 complex apparatus; found linear relation between pupil diameter and (1) accommodation, (2) accommo- dative vergence, and (3) fusional convergence; latter is minor in some subjects; possibly an influence of psychic awareness of nearness; under normal condi- tions pupil reflex is essentially elicited by accommo- dation 19bOa Marg & Morgan * psychic proximity factor influenced pupil in only 1 of 10 subjects; pupil reaction elicited by accommodation; vergence elicited by accommodation, does not occur alone under normal conditions 1950b Marg & Morgan * - no pupillary effects of fusion found, independent of converge nce -accommodation
73 TABLE 9 (Cont'd) Experiments on Man x I c W (0 cfl 01 1 u 1 X, 1 Â£ â¢ jj U Year Author j cu r- I 1 Findings and Conclusions 1951 Renard & Masonnet- * * * pupil contraction is not reflex but associated move- Naux ment (with convergence) 1956 Lowenstein * * obtained isolated accommodation, contraction internal rectus, miosis by electrical stimulation of 3rd nerve; case with present light reflex, absent near vision contraction OD, absent light reflex, present near vision contraction OS 1956, 60a Shakhnovitch * , records eye movements and pupil together; conver- 61 gence movement is complete before end of latent period for pupil 1958.59 Jampel * studied supranuclear control of ocular movements in monkey 1961 Alpern, Mason, & * complex apparatus; found linear relation between Jardinico accommodative vergence and pupil, curvilinear rela- tion between accommodation and pupil; discrepancy thought to be due to limitations of amplitude of ac- commodation imposed by lens 1961 Samojloff, Sokolova, * pupillary reaction symmetrical when only one eye & Shakhnovitch converges (uncentered target) 1924-65 Lowenstein * diverse cases: isolated accommodation paralysis. Adie's syndrome with or without tonic accommoda- tion, loss of pupil contraction with preserved accom- modation and /or convergence report of a. finding by Miiller (1854) is still soberly quoted today (Smythe, 1958) to the effect that in the dog, in contrast to other mammals, the pupil dilates upon near vision and contracts upon far vision. But the visual target provided for the dog was a piece of meat or sausage. Trautvetter (1866), who repeated the experiment with similar results, described how the dog, having been starved for two days, "followed the sausage with the most desirous of glances" 10. Obviously, the dog must have been under more than visual stimulation as the deliciously fragrant "".. .Ich griff sogar zu kiinstlichen Mitteln, um die Thiere zu zwingen, ihre Augen fur die Nahe und Ferae einzustellen, und liess zu diesem Zwecke einen jungen Hund und eine Katze zwei Tage hungern; dann band ich den Hund auf das Brett und indem ich selbst die Linsenbilder beo- bachtete, liess ich einen Gehulfen nach Kommando ein Stuck Wurst der Nase des Hundes nahern und dann wieder von derselben entfernen . . . Das Thier verfolgte die Wurst wahrend der ganzen Zeit des Versuches mit den lusternsten Blicken. . .Bei der Katze, der zum Accommodations- gegenstand eine lebende Maus diente, misslang der Versuch, da sie, wohl ihrer tiickischen Natur wegen, sehr nachlassig accommodierte. . . "
74 TABLE 10 More Elaborate Experimental Designs Year Author Chief Purpose of Design 1869 von Arlt time pupillary light reflex characteristics 1880 Angelucci & Aubert time movements of pupil and Purkinje images 1893 Sachs test color sensitivity 1894 Greeff & Du Bois-Reymond relate pupil size to stimulus luminance 1897 Garten record pupillary dilation after withdrawal of light 1900b,c Abelsdorff test color sensitivity 1900 Vervoort test relations between pupil, accommodation, and convergence 1903 Friberger measure time characteristics of pupillary movements 1903 Schafer test color sensitivity 1904 Abelsdorff & Feilchenfeld test pupil response to stimulus luminance and area 1905 Basier test color sensitivity 1907 Hess explore retinal areas (pupil perimeter) 1907 Polimanti test color sensitivity 1907 Schlesinger fine pupillary thresholds to white or colored stimuli 1908a Hess test color sensitivity in birds 1908b Hess find response to alternate stimulation of different retinal areas 1910 Hess test color sensitivity in animals 1913 Schlesinger objective perimetry (peri-pupillometer) 1914-16 Hess test difference threshold (white or colored light) 1918 Blanchard study retinal sensitivity (brightness, contrast, etc.) 1919 Engelking test pupillary threshold 1918,20 Reeves study rate, extent of pupil contraction and dilation 1923 Laurens test spectral sensitivity (photopic and scotopic) 1923 Cradle & Eisendraht study speed of pupillary contraction to light 1926 Holladay study reactions to glaring light 1926-65 Lowenstein study effects of stimulus luminance, duration, frequency 1929 Barbieri perimetry (pupil perimeter with colors or white light) 1929,30 Stiles study effect of glare on brightness difference threshold
75 TABLE 10 (Cont'd) Year Author Chief Purpose of Design 1932 Cradle & Ackerman study speed of pupillary redilation after light stimuli 1933 Ferree, Rand & Harris determine effectiveness of very large adapting fields 1933-35 Machemer study dynamics of pupillary reflexes to bright light 1934 Biffis test sensitivity of different retinal areas 1934 Luckiesh & Moss determine effects of stimulus area and luminance 1936-37 Crawford determine effects of stimulus area and luminance 1938,43 Kappauf study brightness sensitivity in cat 1938 Talbot find influence of stimulus intensity, area, duration 1939 Brown & Page record pupillary dilation after withdrawal of light 1940 Frydrychovicz & Harms objective perimetry (pupil perimeter) 1939-40 Hecht & Pirenne determine spectral sensitivity in owl 1942a Bartley record reactions to light flashes at different rates 1942 Wagman & Gulberg determine pupillary spectral sensitivity (scotopic) 1942 Wagman & Nathanson study effect of stimulus luminance in man and rabbit 1943 Bartley test stimulus luminance, area, binocular interaction 1945 Fry study relation of pupil, accommodation, convergence 1948 Barany & Hallden study retinal rivalry 1948 Flamant test effect of stimulus luminance for very large field 1948 Spring & Stiles test effect of light entering the eye via different areas of the pupil 1949 De Launay study effect of stimulus luminance 1949-54 Harms objective perimetry (pupil perimeter) 1949 Knoll relate pupil changes, accommodation, convergence 1949,50 Marg & Morgan test relations of pupil, accommodation, and convergence 1950 Campbell & White side record pupillary oscillations in steady light 1951-54 Cuppers relate pupillary reactions to retinal sensitivity 1953 Alpern & Benson show directional sensitivity of pupil receptors 1953 Fry & Allen study pupil responses to light flashes of different luminance, area, duration, retinal location 1954 Young & Biersdorf record pupillary reactions to light and darkness 1954 DuBois-Poulsen & Loisillier explore retinal areas (dark-adapted eye)
76 TABLE 10 (Cont'd) Year Author Chief Purpose of Design 1955,56 Schweitzer study pupil threshold of dark-adapted eye (various areas, durations, colors, retinal positions of stimuli) 1956 Fugate & Fry study pupillary movements to glaring light 1956 Hopkinson study pupillary reactions to glaring light 1956 Van der Tweel study effects of stimulus luminance, rate, wave form 1957 Bleichert & Wagner record pupil responses to sinusoidal light stimuli 1957 Shakhnovitch record pupillary and eye movements during reactions to light and to near vision 1957 Stark & Sherman servo-analysis of pupillary reactions (frequency) 1957 Stegemann servo-analysis (sinusoidal light stimuli) 1958 Samojloff & Shakhnovitch explore different retinal areas (objective perimetry) 1958-60 Shakhnovitch record pupillary and eye movements during near vision 1958 Stark & Campbell servo-analysis of pupil oscillations in steady light 1958 Stark & Cornsweet servo-analysis of pupil oscillations in steady light 1959 Alpern, Kitai, & Isaacson study dark-adaptation of pupillomotor receptors 1959 Lowenstein & Loewenfeld study pupil threshold (light- and dark-adapted eye) 1959 Samojloff objective perimetry 1959 Shakhnovitch study color sensitivity in cat 1959 Stark servo-analysis of pupil reactions (stimulus frequency) 1959 Stark & Baker servo-analysis of pupillary movements in steady light 1960b Shakhnovitch objective perimetry 1961 Alpern, Mason, & Jardinico test relations between pupil, accommodation, convergence 1961 Samojloff, Sokolova, & Shakhnovitch record eye movements and pupillary changes during near vision and other eye movements 1962a Alpern & Campbell determine photopic pupillary spectral sensitivity 1962b Alpern & Campbell study pupil during dark-adaptation after bright light 1962 Bouma determine pupillary spectral sensitivity (photopic) 1963 Burke relate visual and pupillary thresholds 1963 Feinberg & Podolak determine latent periods of light reflex 1964 Lowenstein, Kawabata, & Loewenfeld relate visual and pupil thresholds, visual flicker fusion, and pupillary incremental behavior