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The effect of temporal, frontal and parietal lobe ablations on the maze behavior of cats Griffiths, Roy Scott 1954

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THE EFFECT OF TEMPORAL, FRONTAL AND PARIETAL LOBE ABLATIONS ON. THE MAZE BEHAVIOR OF GATS, by ROY. SCOTT. GRIFFITHS A THESIS SUBMITTED IF. PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of Neurological Research We accept this thesis as conforming to the standard required from candidates for the degree of MASTER OF SCIENCE. Members of the Department of Neurological Research THE UNIVERSITY OF BRITISH.COLUMBIA October, 1°54. ABSTRACT In the present investigation the effect on brightness discrimi-nation of ablations of various parts of the cerebral cortex of cats has been examined. The background for this project lay in the work of Kluver and Bucy (1939) who observed "psychic blindness0 in monkeys with bilat-eral temporal ablations and that of Chow, Blum and Blum (1951) and Rlopelle, Alper, Strong and Ades (1953) who found no such loss of simple visual properties from similar ablations in monkeys. Since i t is not possible to subject the cat to the complex experimental tests of the monkey, and since a rigorous control of the stimulus situation was desired, the present investigation deals with brightness discrimination only. It was found that bilateral ablations of the cerebral cortex had a deleterious effect on bright-ness discrimination. The specific test did not differentiate among the effects of ablations of the frontal, temporal and parietal associa-tion areas although the total post-operative behavior of the animals easily indicated the specific site of the lesions. ACKNOWLEDGMENT This problem was conducted under the supervision of Dr. Margaret A. Kennard of the Department of Neurological Research, University of British Columbia and I wish to express my sincere appreciation to her for much helpful advice and direction. I am indebted to Dr. M. Sam Rabinovitch of the Department of Neurological Research and to Dr. Douglas Kenny of the Department of Psychology of the University of British Columbia for their many helpful suggestions and criticisms. Appreciation is also due to the other members of the Department of Neurological Research, University of British Columbia, who have helped in a great many ways - and thanks to my wife for her unending en-couragement and patience. In addition I would like to thank Dr. A. J. Wood of the Department of Animal Nutrition of the University of British Columbia for hia co-operation and for the use of his facilities in providing an experimental room. None of these persons is respon-sible, however, for any faults of this study. I am especially grateful to Dr. tfm. C. Gibson, Chairman of the Department of Neurological Research, University of British Columbia, for his generous assistance and kindness in making i t possible for me to carry out this work in his department while I was a recipient of a research studentship under the Federal Mental Health grants from Ottawa. TABLE OF. CONTENTS Page Abstract Introduction IL A review of the approaches to the study of cerebral functioning. A. Cytoarchltectonic 2 Bv Histologic 4 1. Frontal areas 4 2. Parietal areas 4 5. Temporal areas 5 C Electrophysiologic 5. 1. Frontal associations areas 65 2. Parietal association; areas 6 5. Temporal association areas 11 EL. BehavibralL 8; 1..Frontal areas 8 2. Parleto- temporal areas %• E. Clinic 15 L. Frontal areas 15 2. Parietal areas 14 5. TJemporal areas 15 F. The Problem 16 Materials and Procedure is-A. Place ia B.. Subjects 18 1. Housing 18 2. Pood 19 a. Apparatus 19 D. Techniques 22 1. Training 22 2. Criteria of testing 25 5. Statistical 26 4. Surgical 27 Results 29 A. Brightness discrimination data 50 Table of Contents (Continued) Page B. Behavioral observations 35 1. Frontal group 35 2. Parietal group 38 3. Temporal group 38 4. Additional ablations 40 5. Additional training 41 Discussion 42 A. Frontal group 42 B. Parietal group 44 0. Temporal group 45 D. Additional ablations 46 Summary 48 Bibliography 51 Appendices A. Statistical Tables Table I! Trial order to avoid scores due to position: habits, simple alternation and double alter-nation (Modified from Gellerman, 1933). Table II Criteria for determining length of run re-quired for statistical significance with different numbers of total trials (After Grant, 1947). Table III: Discrimination test results Primary parietal: a, h, c Primary temporal: d, e, f, g, h, i Primary frontal: j , k, 1, m, q Included in pre-operation training: n, o., p, r. Table W Mann-Whitney U-test for n - 3, 4, 5, 6, 7. Table V Critical values for r for Sign Test (From Dixon and Massey, 1951)* B.. Cortical maps and ablations Plate I Map of Monkey brain (After Brodmann, 1909) Plate II Map of Cat brain (After Garol, 1942c). Plate III Cortical maps Plate 17 Cortical areas assumed for ablations Plate V Maps of ablations. a) Primary parietal lesions Table of Contenta (Continued) b) Primary frontal lesions c) Primary frontal lesions d) Primary temporal lesions e) Primary temporal lesions C. The Apparatus Plate I Plan of the maze. Plate II Top view. Plate III Side view. Plate IV. Waiting cages. 1 IHT310DU0TI0N In the investigation to be reported here, the effect of lesions of the cerebral cortex on the behavior of cats in response to visual stimuli has been observed. In particular, the focus of attention has been directed to the functions of the temporal lobes which have been compared to those of the frontal and parietal areas. Brightness dis-crimination has been selected as a unit of simple visual discrimina-tion although i t is well known that there are other visual qualities -among them are object or pattern discrimination (Morgan, 1951) - which can also be differentiated by the normal cat. Cats were chosen for the experimental animals here for several reasons. They are capable of training in visual discriminations; they have a relatively simple cerebral cortex as compared to that of the monkey; this particular in-r vestigation has not been carried out on this particular species; and finally, cats were, at the time of this investigation, readily avail-able. Although considerable experimental work has been done in the last half century on the various behavioral characteristics of animals, i t is only recently that much interest has been directed to the temporal lobe. Clinical material and the work on monkeys of such investigators as Klftver aad Bucy (1959) and others (Adas and Raab, 1949, Meyer, Harlow and Settlage, 1951, Riopelle, Alper, Strong and Ades, 1955) has shown that the changes in behavior which follow bilateral temporal lobe ablation are related in some degree to visual discrimination. No 2 such relationship has been found between visual function and the patterns of behavior which follow removal of either frontal or parie-tal regions (Chow, Blum and Blum, 1951, Harlow, Davis, Settlage and Meyer, 1952). For purposes of general orientation further material and informa-tion which contribute to the background of this study will be discus-sed under various headings as follows: A. Cytoarchitectonlc. For the present study i t was necessary to select and define the cortical areas which were considered as belonging to the cerebral cortical units to be ablated in the cats. This was done, using in-formation obtained from either anatomical or physiological studies. It will be seen that there is some indefiniteness as to exact bound-aries. The anatomic information was based on the combined agreements and disagreements of a number of investigators (Campbell, 1905, Brod-mann, I9O9, Ariens Kappers, I929), who have mapped the cerebral corti-cal areas cytoarchitecturally, that ia, according to the histological appearance of various cellular, cortical layers. Brodmann (1909), has assigned numbers to the various areas which are those used here (Plate I, Appendix B). The association areas in the frontal, temporal and parietal regions are those which have most recently developed phylogenetically and which, as far as is known, serve to elaborate the 'higher functions' of the adjacent sensory areas. In man, chimp-anzee and monkey, the association areas are relatively large and have been, very probably, overdivided by this cytoarchitectonlc method. In the cat, however, the association areas are relatively small 3 (Poljak, 1927). They are shown in Plate II, Appendix 5, as outlined by Garol (1942c) adapted from the work of Brodmann. The indefinitenese of cyt©architectural boundaries is well i l -lustrated by the work of the following authors. Lashley and Clark (1946) found no basis for division of temporal cortex into Brodmann1s areas 20, 21, 22 on a structural basis. Nor could they find any clear line of demarcation between temporal and occipital lobes. They found, studying ateles and macaque monkeys, only general character-istics for frontal, precentral, postcentral, parieto-occipital, temporal, striate and insular areas. Subdivisions of these areas on a basis of fixed criteria was impossible. They are in agreement with Bailey and von Bonin (1951) when they say, "What was needed was to state as objectively as possible those architectural types that can be clearly distinguished, but to refrain from giving quite secondary, obviously unimportant details the same critical value." Bailey and von Bonin state further, °To draw a map on which these three areas are given three different markings - such as dots, cross-hatchings, and broken lines is to create an entirely misleading impression. Useful as such maps are for the description of cortico-cortical connections, they do not translate accurately cytoarchitectonic data." "This is not to deny that such subtle and minute distinctions can be made; i t is, however, to deny that, at least at present, they serve any useful purpose. Coming from subhuman primates we are more impressed, as will appear in the course of this monograph, by the astounding homogeneity of the human cortex than by its divisibility into a host of elemental organs as Brodmann would 4 have i t . 0 B. Histologic. The cerebral cortex is in close connections with deeper structures, particularly the thalamus. By degeneration studies the frontal, parietal and temporal association areas can be related to the particular nuclei to which each area is directly connected. Using this approach the f o l -lowing relationships were noted. 1. Frontal Areas. The frontal areas project to both the anterior and the dorso-medial nuclei of the thalamus. The anterior thalamic nuclei also re-ceive a large projection from the mamillary bodies via the tract of Vicq D'Azyr and project in turn to the anterior cingulate gyrus (Le Gros Clark, 194s). The orbito-frontal cortex has been found to be the pro-jection areas for the dorsomedial nuclei of the cat (Rose and Woolsey, 1948). Le Gros Clark and Boggon (1955) demonstrated a connection through the periventricular system between the hypothalamus and the dorsomedial nucleus. 2. Parietal Areas. For the most part the posterolateral nuclei of the thalamus and the pulvinar project fibres to the parietal associative cortex. The more caudal cortex receives ita fibres from the pulvinar nuclei (Minkowski, 1951, Chow, 195°, Walker, 1958). There are U-shaped, short association fibres passing in a complicated fashion so as to link up a l l neighboring areas of the homolateral parietal cortex. There are also commissural fibres which, taking origin in one part of the parietal cortex, run trans-veraly across the middle third of the corpus callosum and terminate in a corresponding point in the opposite hemisphere. But Critchley (1953) states, "The exact distribution and conformation of such cortico-cortical fibres are not known for certain. 0 3. Temporal Areas. According to Walker (1938) the temporal lobe has a paucity of connections with the thalamus. Extensive degeneration within the pulvinar i s seen only when the cortex of the posterior part of this temporal area i s damaged. Mettler (1935) found the precentral and postcentral cortices to be poorly associated with the temporal cortex. In reporting the anatomical changes secondary to temporal lobectomies of the monkey, Bucy and Klftver (1940) found very similar connections. In addition they found degeneration in temporo-frontal fibres, and in two large groups of fibres, one of which passed upward and backward to the corpus callosum and the other to the posterior cingulum. G. Electrophysiologic. By the technique of physiological neuronography originated by Dueser de Barenne (1910), c e l l activity i s potentiated by strychnine applied to the cortex. Using this method, DusBer de Barenne states in his a r t i c l e with Garol and McCulloch (1942) that he was able with experiments carried out in 1911 to show that strychnine when applied locally acted s t r i c t l y locally; that i t excited only where there are synapses on nerve c e l l s ; that i t produced disturbances which travel only in the direction of normal conduction, not antidromically; and f i n a l l y that these disturbances were diminished, dispersed and de-layed when they reached the primary axonal terminations. The cur-6 rent hypothesis as to the mechanism whereby this is brought about, ia that strychnine does not increase the activity of a cortical neurone, but rather synchronizes the cellular activity in the six cortical layers leading to the production of "strychnine spikes" (Dusser de Barenne, Marshall, Kima and Stone, 1941). Two other electrical phenom-ena, that of evoked potentials, which follow a single pulse stimulus, and that of after discharge, which follows a repetitive electrical stimulus, can be used separately or in combination with the strychnine method. Some of the findings for the three association areas, as found by the above procedures are indicated below. 1. Frontal Association Areas. Frontal association areas include Brodmann's areas 9, 10, 11 and 12 (Plate I, Appendix B). Ward and McCulloch (1947) were able to demonstrate direct connections between the orbital regions of the frontal lobe and certain hypothalamic nuclei. Sachs, Brendler and Fulton (1949) found strychnine applied to orbital area 15 of one side activated the putamen and caudate nuclei of both hemispheres. Bailey, von Bonin, Garol and McCulloch (I94jb) also using physiological neuronography have shown long cortical association tracts between area 47, that is, orbital surface and the tip of the temporal lobe (area 58)• The above mentioned workers found in addition, from stimulation of area 8 known as the frontal eye field, well-defined strychnine spikes were produced which spread to the parastriate area of the ipeilateral and contralateral hemisphere. 2. Parietal Association Areas. For working purposes in this experiment, the parietal as-7 aociation areas are considered to include Brodmann's areas 5 and 7 (Plate I, Appendix B). Dusser de Barenne, Garol and McCulloch (1941) found, from neuronographic studies of the monkey, connections between area 7 and the f i r s t temporal sulcus. In addition, they demonstrated, from strychninization of area 5* f i r i n g within i t s e l f and f i r i n g of areas 7, 2s, 1 and 4. Strychninization of area 7 produced local f i r i n g within area 7 and f i r i n g in areas 5» 2s 1 and 4. Strychnine applied to area 19s produced spiking within this area only. Bailey, von Bonin, Garol and McCulloch (1942b) demonstrated strychnine spikes in the temporal lobe of the monkey from stimulation of area 18. Gen-erally stimulation of the more inferior portion of area 18 propagated further toward the tip of the temporal lobe, that i s , into the central portion of the upper l i p of the second temporal sulcus but never to the extreme t i p . 3. Temporal Association Areas. Brodmann's parcellation of these association areas i s 28, 20, 21, 22. Other workers (Bailey, von Bonin, Garol and McCulloch, 1942a) include area 38 which surrounds the temporal t i p . The cortico-cortical fibre connections have been mentioned in the f i r s t two sub-divisions of this section. It should be noted that the connections of these areas are in the order of frontal-parietal, frontal-temporal and parietal-temporal (Bailey, von Bonin, Garol and McCulloch, 1943b). Reverse connections have not yet been demonstrated. Commissural fibres connections the two temporal association areas have their course through the anterior commissure and psalterium (Marsan and S t o l l , 1951)* S t o l l , Marsan, and Jasper (1951) i n an elaborate study using 8 a l l three electrophysiological methods, demonstrated connections of the temporal tip with numerous subcortical nuclei in the cat. The findings of these workers, particularly the connections of the temporal tip with the caudate nuclei, hypothalamus and pulvinar, would indicate an anatomical link between a l l three association areas. D. Behavioral. Psychological examination of the behavior of animals has been carried out in a number of different ways. The most common method is that of instrumental conditioning in which the animal is trained to perform some task using its natural preference and motivation. Various portions of the nervous system may then be removed and the results on its trained behavior noted. The early investigators, Ferrier (1886) and Franz (1°07) performed ablations on 'normal' animals and then reported observations as to general changes in behavior. Yerkes and Watson (1911) and later Lashley (I929) began to quantify the observa-tions limiting their fields to specific test situations. 1. Frontal Areas. The function of the frontal areas has been defined by a number of investigators. Jacobsen ( 1 9 5 6 ) discovered, from pre- and post-operative comparisons on monkeys with ablations of the frontal association areas, what is now considered to be the classical symptom of these animals, that is, loss of ability to solve problems involving delayed response. He failed to find any change in the monkeys' ability to solve the problem box, or visual discriminations. These results have been confirmed by otherB and elaborated on, mainly by Harlow and his associates ( I 9 5 I ) . In Harlow's opinion frontal lesions impair the 9 solution by the rhesus of a l l difficult and complex problems. Such difficult tasks are the oddity problems and double alternation (Harlow. 1955)• Although there has been l i t t l e work done with chimp-anzees, i t would appear that there is a species difference as i t has been found that frontal lobectomies have l i t t l e deleterious effect on the delayed response in these animals (Ackerly, 1950). Extending this work to humans, the Columbia-G-reystone Associates (19^9) studied the results of a battery of thirty-five psycholbgical tests on nineteen psychotic patientB who had had bilateral partial ablation (topectomy and gyrectomy) of the frontal cortex. They found °no real or perma-nent impairment of mental function brought about by the operation, which could be demonstrated in any way by our exhaustive psychologic test battery 0. This may point out the inadequacies of the tests used. It may also indicate that there is an added uncontrolled factor of in-complete supervision over the past learning histories. This assumes that i t is possible for the individual to develop, in his lifetime, several alternative ways to attack the problems of a test that is sup-posed to measure a function. Lastly, there is the possibility of a species difference which was mentioned above by Ackerly to be existent in the subprimates. 2. Parieto-temporal Areas. The parietal and temporal cortices are being combined in this section since l i t t l e work has been done with regard to vision on the parietal association area. Ruch, Fulton and German (1958) found the parietal lobe involved in fine weight discrimination, as such discrimination was abolished with ablations of that area. It 1 0 has been found by Harlow, Meyer and Settlage ( I 9 5 I ) , Harlow, Davis, Settlage and Meyer (1952) that ablation of parietal association areas of monkeys does not have any effect on delayed response reactions, yet produces a great deficit in such complex tests as oddity problems. Chow, Blum and Blum (1951) from their experiments on monkeys, are in agreement with Lashley that lesions confined mostly within the pre-occipital area f a i l to cause any deficit in visual function. Further-more, they state, "The temporal focus of visual function appears to be confined to the cortex. Degree of damage to the thalamic relay and aesociative nuclei ia not correlated with visual disabilities. 1 1 This is an interesting finding when compared with the early work of Kluver and Bucy (1939) where a large portion of the underlying structures such as hippocampi, amygdalae, t a i l of caudate and uncus were removed or damaged. Poirier (1951) who also ablated these subcortical nuclei, as well as cortical structures, was not able to find any symptom of "psychic blindness" as described by Kluver and Bucy. From the many experiments of Allen (1940, 1941) i t is fairly well established that the hippocampi, amygdalae and pyriform areas are not in any way associated with conditioned olfactory responses. This gives credence to the histological findings of Brodal (1947) that the hippocampus can be considered a cortical structure. Certainly from its connections with the cingulum, i t would appear to be involved in behavioral changes (Adey, 1951). Bard and Mountcastle (1945) produced rage in cats with ablations that removed or damaged the amygdaloid com-plex and much of the cortex of the pyriform lobe. Spiegel, Miller and Oppenheimer (1940) produced savage rage outbursts in the same species 11 with electrolytic lesions in the amygdaloid nuclei, but this iB in marked contrast to the changes observed in monkeys by Klftver and Bucy (1959) and by Poirier (1951) who found greatly reduced anger and fear reactions after ablation of these nuclei. Yet presumably there is, as Papez (1957) suggests, an anatomical circuit involving the temporal cortex, hippocampus, thalamus and cingulum in the elabv orating and regulating functions of the central emotional state. A number of psychologists consider changes in emotional state to have influences on visual perception. Ades (1946) achieved similar results to Chow and Lashley with lesions of the prestriate areas, 18 and 19. He found the learning curves in his results to be the same pre- and post-operatively. Prom this he concludes "areas 18, 19 are essentially involved in i n i t i a l development of particular patterns of visual interpretation or associa-tion, since habits disappear because of the operation, but return with retraining". He does not speculate on the "unknown masses of visual matrix" which are concerned in the mechanisms involved. In an extension of this experiment, Ades and Raab (1949) endeavored to determine the effects in monkeys of additional ablation of the temp-oral lobe on learned visual discriminations. These workers did not find "psychic blindness" although they noticed a tendency to nose a l l food. They found, as Ades had before (1946), that visual discrimina-tion can be learned again after bilateral ablation of areas 18, 19. They also found that bilateral ablation.of the temporal lobes alone had no effect whatever on the relearning of this habit. But i f areas 18, 19 were ablated after the temporal lobes, then no amount 12 of training would re-establish the habit. From this Ades and Raab deduce that the temporal cortex "is not an essential link in the visual chain through whose activity the visual discrimination is originally learned in the intact animal; neither does its absence impose an appre-ciable delay in the i n i t i a l learning. It is equally evident that the temporal cortex (or a part of it) becomes essential to the process in the absence of areas 18, 19°. There is evidence of a different type of loss of intellectual function following lesions of the frontal and parietal association areas from the experiments of Chow, Blum and Blum (1951) on monkeys in which they add bilateral lesions of the frontal association areas to bi-lateral destruction of the posterior association areas. The posterior removal produced a syndrome including the loss of, and retardation in, relearning habits of visual discrimination. The addition of frontal lesions affected various components of this syndrome differently, but i t is of interest here that there was no effect on visual discrimination, although delayed reaction responses were further impaired. Harlow mentions in his review (1953) unpublished results similar to those of Chow and his co-workers. Bilateral frontal lesions were produced in two monkeys with bilateral posterior association area lesions of two years' standing, and bilateral destruction of the poaterior association areas waa effected in two monkeys with long established frontal associa-tion area lobectomies. The new, posterior lesions produced drastic impairment in difficult discrimination tests but produced no additional deficit in delayed response, and the new frontal ablations resulted in serious impairment of the delayed response performance but no additional 15 deficit in visual discrimination problems. It is apparent, then, that there is, for the monkey at least, good evidence of cortical localization of function in the association areas. S t i l l the evidence is by no means unequivocal as there are studies which uphold the concept of equipotentiality (Levine, 1952) and even a study on the rat which is felt to disprove this concept of equipotentiality (Pickett, 1952). According to Harlow (1955) "the theory of cortical equipotentiality would require that sensory areas mediate intellectual as well as receptive functions". But the separa-tion of sensory, perceptual, and intellectual loss in any definite way is difficult, for these functions, which may not be discrete, interact in very complex ways. With the truly phenomenal degree of recovery of function from cortical lesions that we now know is possible with training, i t appears doubtful according to Harlow (1955) that a station-ary lesion will produce a permanent loss of one specific "psychological process". E. Clinic. In this section the main concern will be with epilepsy and the information i t has contributed to understanding the function of the frontal, parietal and temporal areas of the brain. 1. Frontal Areas. For the most part epileptic seizures that are associated with foci in the frontal association cortex are not preceded by aurae, but rather there is an immediate loss of consciousness with adversive movements following i f the frontal eye fields are involved. Either symptom may appear separately (Penfield and Uristiansen, 1951)• Petit 1* mal attacks, although they are characterized by electroencephalograph^ changes fi r s t in the mid-frontal lobes, are thought by some (Jasper and Fortuyn, 19^ 7) to have origin in Borne subcortical centre because of the speed with which the discharge spreads to involve the occipital regions. Furthermore, a major seizure in petit mal patients may begin with a crescendo-like increase in the voltage of the alpha rhythm. It is possible, then, that there is a connection here with perception, including visual, through an influence on attention. 2. Parietal Areas. In patients with epileptogenic foci in the anterolateral, parietal association areas, Penfield and Kristiansen found vestibular disturbances (dizziness, rotatory sensations and giddiness) to be most common. Excluding the simple sensory components of an epileptic seizure in the parietal lobe since such sensations arise from pathology in the anterior parietal area, we find, from the interparietal areas, as described by Critchley (1953) a syndrome of complex dysesthesia; "Complicated phenomena relating to the body-image may develop belong-ing to the category of what Bonnier termed 'paraschematia'. Thus a limb may seem to shorten at the expense of the proximal portion, so that the hand (or foot) which may also seem to be in violent involun-tary activity, may appear to become telescoped up towards the trunk." Visual disturbances from this area are associated with penetrating lesions. Purely visual disturbances such as object displacement and inability to recognize colors were found by Hoff and Kamin ( 1 9 5 0 ) . Olson and Ruby (1941) found a vertiginous symptom associated with 15 lesions in this region. 5. Temporal Areas. Of particular interest to this study is psychomotor epi-lepsy which, in recent years, has been widely investigated. As op-posed to frontal lobe seizures, where there are no aurae, the temporal lobe seizures are very often preceded by vestibular, auditory, or visual aurae. People afflicted with this type of epilepsy may experi-ence what is termed the deja. vu phenomenon. In this experience the individual in a new situation believes i t to be some familiar experience that he has had before, with either visual or some other sensation such as smell or taste. Psychomotor epilepsy is characterized electro-encephalographically, say Gibbs, Gibbs and Fuster (1948), by a focal, spike, seizure discharge in the anterior temporal region with a fre-quency of 4 to 6 cycles per second. The clinical symptoms are confusion and amnesia. There is usually no loss of consciousness, movements appear purposeful, yet poorly co-ordinated, and more or less automatic. Prom their studies of the cortical stimulation of epileptic brains, Penfield and Boldrey (1959) found that habitual seizures have a conditioning effect upon neuronal pathways which may establish a complicated neuronal pattern so functionally organized that i t may be produced by electrical stimulation, although no such complicated res-ponses could be elicited from a normal brain. The fact that these temporal lobe seizures are characterized by alterations in perceptual vision (during the seizure the patient is aware of his surroundings but there is a sudden change in his interpretation of visual percep-tions) or by hallucinations (he has the illusion of dreaming and being 16 afraid of the dream, yet is aware of his surroundings) led Penfield and Jasper (19^7) to hypothesize that since the hallucinations were made up of remembered fragments, the acquired pattern of neuronal connections in the temporal lobe actually constituted the record of such memory. Jasper, Pertuisset and Flanigin (1951) from investiga-tions of electroencephalograms and cortical electrograms, classify patients with temporal lobe epilepsy into four groups; those with 1) epileptoform discharge in one lobe with no conduction of discharge to the other side, 2) a clearly localizable focus in one lobe, but with intermittent low voltage waves transmitted to the temporal lobe of the opposite side, 5) epileptic discharge of equal voltage appearing synchronously from the two lobes or 4) localized temporal discharge shifting from one lobe to the other. From their evidence they were led to believe that the first three groups were primarily cortical epileptogenic foci, while the last group had a primary focus in some deeper structure. This gave rise to the work of Marsan and Stoll (1951). F. The Problem. From the experiments of Kl&ver and Bucy (1929) with their observa-tions of "psychic blindness" and the findings of clinicians with res-pect to alterations in the sensory perceptions of individuals with psychomotor epilepsy, i t was decided to study the effects on behavior of lesions of the temporal and frontal association areas in order to determine whether such perceptual changes could be reproduced and quantified for the cat. The ablations of the frontal association areas were included in this study in an effort to ascertain whether or not the changes in affective behavior produced by lesions in these areas 17 would affect visual perceptions. The parietal areas were used for a control ablation. Several investigators (Chow, Blum and Blum, 1951, Meyer, Harlow and Settlage, 1951* Ades and Raab, 1949) have examined this field with monkeys and i t was thought that their lack of posi-tive findings in visual tests might be due to the complexity of the subprimates1 behavior. With this in mind, the present study was under-taken in the hope that the much less complex behavior of the cats could be more easily analyzed and give a better indication whether or not there is functional localization of visual discrimination in the areas studied. 18 MATERIAL AND METHOD A. Place. The conditioning experiments were carried out in the Department of Animal Husbandry and the surgical procedures were performed in the Department of Neurological Research, University of British Columbia, from October, 1952, to August, 1955. The conditioning was carried out in a quiet, darkened room. Although not completely soundproof, there were no disturbances which appeared to affect the cats' behavior. This experimental room was some distance from the room in which the cats were quartered and the animals were transported in a carrying cage 45 x 25 x 28 cms. A l l animals were kept under uniform conditions throughout the experiment, that is, the building was air-conditioned and thermostatically controlled and they were maintained on a constant diet. B. Subjects. 1. Housing. The subjects were eighteen adult cats (sixteen males and two females), none of which had previously been trained or used for any experimental work. After being quarantined for two to six weeks to exclude infections, they were placed in individual cages measuring 55 x 70 x 47.5 cms. for a further period of two to five weeks to ac-custom them to the living quarters, and trainer, and to standardize the hours of feeding and the amount of food. After the training pro-gram began the cat was never again fed in its cage, although water was 19 there at a l l timet}. 2. Food. The general care and diet of the experimental animals has been standardized by Dr. A.J. Wood of the Animal Husbandry Department. The diet includes: 2 parts fresh frozen meat 2 parts fresh frozen fish 1 part fresh frozen liver 6 parts #10 mash; @ 4 parts water This mixture served as their entire diet and was the con-stituent of the rewards. The daily ration averaged 500-500 grams de-pending upon the size of the cat. The rewards were l i t t l e balls of this mixture weighing 4 grams. GJ. Apparatus. The apparatus used for the experiment was a Y-maze consisting of three sections; a starting box opening into a V-shaped alleyway with each limb ending in a feed box. The starting box was an angle-iron and wire cage 55 c m s * long and 20 c m a « high. Two plywood partitions, made of the same wood as the maze (2-ply* painted a flat ivory color) were placed to give a measurement of 50 cms. wide ( a l l sizes mentioned are inside measurements) at the front and narrowing to 22.2 cms. at © Ration #10 includes as part of 1000 Ground wheat 450 Ground oats 40 Wheat bran 45 Meat 75 Fish meal (70#). 50 Soybean meal 75 Wheat germ meal 75 Liver meal 50 Powdered milk 50 Bone meal 10 Dried yeast 10 Dried grass 5 Apple pomace 20 Salt 5 Beet pulp 20 20 the maze side (Plate I, Appendix C). This box was made 'light tight' with a metal sheet for the floor and a quarter round canopy over the top that waB 25 cms. above the wire top of the starting box at the front side and curved down to meet the top of the box on the maze side (PlateB II & III, Appendix C). The entrance gate to the starting box was 20 cms. wide and 50 cms. high and the maze gate was 16 cms. wide and JO cms. high. The front of the starting box was covered with a plywood shield 112 cms. wide and 75 C S L S» high to hide the operator from the cats' view. Two 1 cm. holes were made on either side of the entrance gate so that the operator could view the cat in the starting box and along the en-tire alley into the feed box. On this screen to the left of the entrance gate was a bare 6.8 volt light bulb providing light for the observer to record his results. The alleyways projected to the right and left 50°' from the perpendicular to the maze gate and were 120 cms. long x 25 cms. wide x 52.5 cms. high. The top of the alleys was covered with a 6 mm. square wire mesh. Over the choice point, or crutch of the Y, was placed a cover in order to increase the differentiation between the light and dark alleys by stopping the light that came from the light source for the experimental room. This light was a 7"§- watt bulb covered by an inverted, conical, opaque shade which directed the light to the ceiling and thereby gave a very subdued and diffuse light. The illumination on the floor of the alleyways and over the table on which the maze stood was equal to 0.5 foot candles. This light was placed directly over the starting box 150 cms. above the floor of the maze and 20 cms. below the light grey ceiling. With the light in this position i t afforded equal_illumination to both alleys and in no way became an 21 extra-naze cue which the cat had to differentiate in the i n i t i a l learning trials. Finally, the feed box which contained a 15 cms. x 5 cms. deep kidney dish, for holding the reward, was 45 cms. wide x 55*5 cms. deep x 5 2 . 5 high. There was a small gate which lifted up at the rear of the feed box and through which the rewards were changed from tr i a l to t r i a l . This gate measured 20 cms. wide x 15 cms. high. As with the rest of the maze these feed boxes were covered with 6 mm. square wire mesh. The maze was lighted from a 6 volt, 5 cell, wet battery. There was a light bulb in the starting box which was over the centre of the entrance gate and 5 7 . 5 cms. above the floor of the box and gave an illumination of 1 .05 foot candles. This light was controlled from a toggle switch at the left side of the entrance gate. At the far end of each alley, just in front of the gate to the feed boxes and projecting down into the alley were located the lights for the visual cues. Rather than present a point of light, such as obtained from using a 6 . 8 volt bulb alone, i t was decided to increase the size of the light source. Two white, plastic, baby's rattles were obtained and the frustum of conical shape with parabolic sides was placed over each bulb. At the base, the frustum measured 5»9 cms. in diameter and projected into the alley 4 . 4 cms. ThiB light source gave an Illumination of 0 . 6 foot candles for the light alley. These lights were operated from a commercial, three pole, mercury switch at the right of the entrance gate. The mercury switch was absolutely silent and thereby afforded no auditory cues to assist the cat in making its discrimination. The various parts of the maze were separated by 1.5 mm thick galvanized iron gates (Plate I, Appendix C). These facilitated the 22 return of the cat to the starting box and also prevented the cat from entering the other alley once i t had made its choice. Over each gate was built a gallows with a pulley on i t through which the string passed from the top of the gate to the plywood screen at the front of the maze. This allowed the operator to raise and lower the gate as the cat progressed through the maze. D. Techniques. 1. Training. In this study instrumental conditioning was chosen as a specific means of analysis of behavior. In order to secure a uniform measure of trials involved in acquisition, the non-correction procedure was selected. This method made a l l trials equal since the animal made a single response on each t r i a l . In contrast, with the correction method, the nature of the trial varied, for the cat might make several responses to the regative stimulus before the trial was terminated. No form of punishment, other than non-reward, was used since punishment would have introduced unknown variables that would be impossible to control. After a period of weeks to allow the cat to become accustomed to the routine, i t was given a series of three pretraining trials. With a minimum of handling throughout the entire experiment, the cat was lifted from its living cage and placed in a carrying cage and trans-ported to the maze. The cat was placed in the starting box and left while the experimenter walked around the maze in a counterclockwise direction to the feed boxes. As quietly as possible an empty kidney dish was placed in the-right feed box and the gate at the back shut; 22 similarly a dish was placed in the left feed box and the gate locked. i The experimenter then continued around to the front of the maze and sat down. For the first pretraining t r i a l the right alley stimulus light was put on. The feed box gates were raised simultaneously, then the alley gates in a similar way, and finally the maze gate. The entire maze was left open and the cat allowed to roam for one half hour. At the end of this time i t was coaxed into the starting box by lowering the gates and withdrawn from the maze. It was then taken from the room and placed in a waiting cage, in the hallway outside the experimental room, for 50 minutes, given its daily ration, allowed 20 minutes to eat i t , and then returned to its living cage. Following the next two pretraining trials, the cat was not fed but left for one hour in the waiting cage before being returned to the living cage. This hunger regime lasted a period of 72 hours. The daily routine consisted of a training period lasting 5 to 20 minutes after which the cat was required to spend 20 minutes in a waiting cage (25 x 45 x 22.5 cms.), then given its daily ration and allowed 20 minutes to eat i t . No cat was fed again until after the next series of trials 24 hours later. The cats were trained and fed six days a week. For the actual training trials, the cat was brought to the experimental room in the carrying cage, then shut in the starting box while the experimenter circled the maze counterclockwise and placed, as quietly as possible, a reward in either the right or the left kidney dish. An equivalent amount of time was spent behind each feed box so that the cat could not distinguish from a difference in the time which 24 dish the reward would be in. It was not possible to be absolutely silent as presumably the cat could hear the experimenter by the noise made by his clothing and rubber-soled shoes as he moved. The motions of picking up a pellet, opening the gate at the back of the feed box, reaching in and placing a reward in the dish (or taking one out), clos-ing the gate, then moving on, were made at the same speed by the experi-menter before each t r i a l . Having returned to the front of the maze, the appropriate light stimulus was switched on and the gates were raised, those of the feed box raised simultaneously, next, the alley gates, and finally the maze gate. The cat's behavior was watched through the ob-servation holes in the plywood screen and once tt had made its choice the gate to the other alley was closed. As the cat left the feed box to return, the gates were closed behind i t . In the early training trials this was found expedient as this procedure accustomed the cat to return quickly to the starting box to await the next t r i a l . From preliminary experimental runs performed in 1952, i t was noted that the cats' behavior in a maze had often Individual peculiar-ities. For instance, many cats have a tendency to turn one way at a point of choice (either right or left) beyond the chance expectancy. Because the maze is a series of such choices, care was taken to have the stimulus presentation sequences arranged BO that the cat with a right-or left-going tendency did not develop such a habit. If a cat at any time started to develop such a preference, the random presentation of the stimulus was abandoned and the cat 'forced' by means of repeatedly placing the reward in the opposite feed box to the cat's preference habit. If the preference was exceedingly strong, the gate to the in-25 correct alley was let down, or at least raised only far enough to prevent the cat from getting under i t and into the incorrect alley. Such 'forced trials' were not counted in the total learning trials for this experiment. 2 . Criteria for Testing. The random order of stimulus sequence used in this study was developed by G-ellerman ( 1 9 5 3 ) (Table, I, Appendix A). It was, however, modified to the extent that each day's number of trials was five instead of ten. It was found from a preliminary experiment that ten trials per day produced no additional learning within the number of trials that had been set, that is, 4 2 9 . This is in keeping with the work of Calvin (1929) who found overcrowding of trials, either having them too close together or too many in one period, to be disadvantageous. It was found a l i t t l e difficult, in the cat, to build up motivation to perform the trials using hunger as the basic drive. Three cats, in fact, had to be dropped from the series because i t was impossible to make them perform even when hungry. A further modification of Gellerman's table was found neces-sary because of the way the trials were broken up to five a day for this experiment. To prevent too many of the ini t i a l trials being to the right or left, i t was decided to run across the table aa numbered in Table I, Appendix A, rather than 1 + 2, next 5 + 6 , next 9 + 1 0 and so on as suggested by Gellerman in his table of stimulus sequences. In any discrimination experiment the scores usually reported are the proportion of correct choices, or the number of trials required to reach criterion. A very adequate table has been compiled by 26 Grant ( 1 9 4 7 ) . From this table i t was decided, using the p = 1/2 column, which applies to this maze problem since the cats had two possible choices, that 429 trials would be sufficient to determine whether or not the cat could learn or relearn the discrimination problem. If i t was not then able to achieve 12 successive, correct choices, i t was assumed i t had not learned the problem. A l l cats used met the criterion requiring, usually, between 200 and 300 trials. In order to use these tables the end of one day and the beginning of the next were counted as part of a run. Once this statistical criterion had been reached, the cat was given a bilateral cerebral ablation and then retrained. 5 . Statistical. The statistical analysis is based on the percentage error scores made by the animals in their response to the light stimuli. With the feed dish 5 cms. deep, i t was not possible for the cat to see whether or not there was a reward there until i t had approached to with-in 30 cms. of the entrance of the feed box. It was not considered an error i f the cat only stepped into the incorrect alley, stopped and retraced its steps to the choice point then went down the correct alley. On this basis the errors made in the entire period of the pre-operative training, divided by the total number of trials required to meet the statistical criterion (Table II, Appendix A), and converted to percent, constituted percentage errors. It was decided to use non-parametric statistics (Moses, 1952) in this study. The advantages of the non-parametric techniques are several. First, whatever the form of the distribution from which the samples have been drawn, a non-parametric test of a specified 27 level actually is that significance level. Secondly, with very small samples there is no other equally appropriate alternative. Finally, the method is usually easier to apply than classical techniques. 4. Surgical. The cats were anesthetized by an intraperitoneal injection of 0.55 cc. nembutal per kilogram of body weight. Rigid aseptic con-ditions were carried out throughout the surgical procedure which followed a ritualistic routine in order to standardize the recovery period as much as possible. Symmetrical, bilateral removals were made at each operation. Through a midline skin incision, the temporalis muscle was reflected and a trephine hole made over the lateral surface of the cortex of one side. The bony defect was enlarged by the use of rongeurs, and the dura then opened. The desired cortical area, frontal, temporal or parietal was then ablated by gentle suction. The procedure was repeated on the second side. The temporalis muscle and scalp were then closed with three layers of silk sutures. During the post-operative recovery period, the animals were carefully watched and, often, hand fed for several days until they were able to be returned to their original cages, when training was resumed. I would like to express my indebtedness to Dr. Margaret A. Kennard who is responsible for the neurosurgery involved in these experi-ments. The actual areas ablated were based on a composite map (Plate IV, Appendix B) which was derived from the varied studies of many workers (Poljak, 1927; Bremer and Dow, 1939; Adrian, 1940; Garol, 1942, a, b, c; Woolsey and Walzl, 1942; Fox, McKinley and Magoun, 1944; Rose and Woolsey, 1948; Harman, 1948) . 29 RESULTS The results presented here include behavioral observations as well as a statistical treatment of the discrimination test. The group of eighteen cats was divided into two, those that had the light cue as the positive stimulus (cats XXVI, XXXVI, XL, XLII, XLVII, LII, LIV, LVI, LVIII), and those that had the light cue as the negative stimulus (cats XXXIII, XXXIV, XXXIX, XLIII, XLVIII, XLVI, XLIX, LV, LIX). This division was made to control any possibility of the cat having some pre-experiment preference for either the light or the dark stimulus. It has been found by Krechevsky (1936) that rats have a tendency to prefer the darker to lighter visual stimuli. In this experiment, the cat was not allowed to respond to such a prefer-ence but was made to respond to the cue that had been predetermined by the group to which i t had been assigned. These two groups were further divided according to the site of the primary cortical ablation. Because of various experimental vicis-situdes only fourteen cats of the original eighteen were carried through to the desired end of the experiment, (two cats (XXXIII, LII) with only temporal ablations became blind, one cat (XLVIII) with a primary frontal ablation developed a permanent obstinate progression, and one cat (LIX) did not complete the in i t i a l learning trials in time to be included in any of the operated groups). Eighteen were used to determine that there was no preference for either the dark or the light alley. The two experi-mental groups were those five having primary frontal lesions (cats XXXVI, 50 XXXIX, XLII, XLVI, XLVII), those six having primary temporal lesions (cats XXVI, XXXIV, XL, XLIII, LV, LVI) and the control group composed of those three with primary parietal ablations (cats XLIX, LIV, LVIII). Initially i t was planned to study the various portions of the frontal lobe. To this end the frontal group was to be divided into orbital, frontal pole and medial sub-groups. However, insufficient numbers were involved (the orbital sub-group consisted of cats XLVII and XLVIII, the frontal pole sub-group contained XXXVI and XXXIX and the medial sub-group was comprised of cats XLII and XLVI) so a l l cats were included, except for cat XLVIII, in the frontal group and statistics were calculated from this group as a whole. Some behavioral differences were noted and these will be mentioned later. A second factor was investigated, that is, the effect of addi-tional secondary frontal and secondary temporal lesions. The secondary frontal lesions were added to three cats (XXVI, XXXIV, LVI) of the group of six that had received primary temporal ablations and the secondary temporal lesions added to the cats with primary frontals (XXXIX, XXXVI, XLII, XLVI, XLVII). The secondary temporal group in-cluded also two cats (XLIX, LIV) with previous parietal lesions. A further control was made of two cats (XL and XLIII) to determine whether or not the anesthetic and the opening of the skull had any effect. These two animals were then given temporal ablations and in-cluded in the primary temporal group. A. Brightness Discrimination Data. The brightness discrimination results are analyzed by non^-parametric statistics. The basis of this analysis is percentage error scores. The 51 tasks at hand were 1) to see whether or not the cats had manifested a dark or light preference in running the maze, 2) to determine whether nor not there were gross differences between the pre- and post-operative scores, 5) to find any differential effects due to the ablation of the various cortical areas. With regard to the fi r s t problem of dark-light preference, the error scores were taken from tr i a l one until the cat had reached the statistical criterion shown in Grant's table (Table II, Appendix A). From the percentages thus computed a Median test (Mood, 1950) w a s applied as follows: Cat No. % Error Cat No. % Error LVI 20.4 . . XLIII 3 6 . 0 XXXVI 22.8 LII 38.4 XLVI 2 7 . 2 XL 39.5 LIV 29.6 XXVI 41.2 XLIX 5 2 . 0 LVIII 41.7 LIX 5 2 . 0 XLII 41.8 XXXIX 32.6 XXXIII 43.8 XLVIII 35.4 XLVI I 46.2 LV 35.9 XXXIV 5 0 . 0 Median = 3 5 . 9 5 Applying the Sign Test, plus for cats' scores above the common median of 35*95 and minus for those below: Dark Stimulus $ Error Sign XLIII XLIX LIX XXXIX XLVI XLVIII XXXIV LV XXXIII 3 6 . 0 3 2 . 0 3 2 . 0 3 2 . 6 2 7 . 2 3 3 . 4 5 0 . 0 3 5 . 9 4 3 . 8 Total +3, - 6 + + Light Stimulus % Error Sign XL 5 9 . 5 • LIV 29.6 LVIII 41.7 + XXXVI 2 2 . 8 XLII 41.8 +• XLVII 46.2 + LII 38.4 + LVI 20.4 XXVI 41.2 + Total + 6, - 5 If we assume there is no difference between the dark stimulus 32 group and the light stimulus group, there should be an equal number from each group above and below the common median, to test this a 2x2 contigency table of frequencies is computed: - + Dark A 6 BJ 3 9 Light a 3 D 6 9 9 9 N; 1 8 Using formula 85& on page 207 of McNemar (19^9), which includes Yates correction for continuity, the following value for Chi Square was obtained; X2 = -892 With df o 1, enter Table 32 on page 242 of Garrett (1947) and P = .45; Therefore, the Null Hypothesis that there is no difference between these two groups could be accepted. A l l the analyses described below are based on error scores de-termined in the following manner. The pre-operative number of trials was divided into thirds, the last third was taken as constituting the cat's behavior once i t had learned the discrimination. These trials were then compared to a similar number of trials immediately post-operative, thus allowing one to determine the effects of cortical ablations on the learned behavior. To determine whether or not there was a difference between a l l cats pre- and post-operatively, a Sign Test (Moses, 1952) was used. Com-paring pre- and post-operative scores of the fourteen cats that under-went cortical ablations, thirteen scores increased post-operatively, one remained the same. Using Table V, Appendix A, the Null Hypothesis could 33 be rejected at better than the . 0 1 confidence level. This indicates that there was a reliable increase in post-operative error scores. The next group of comparisons was made between a l l the primary lesions in the three groups, frontal, temporal and parietal. The object was to determine whether there were any differences in the effects of the three areas. Using the Mann-Whitney Technique (1947), (Table IV, Appendix A), i t was found that there were no significant differences in the pre-operative error scores of the three groups of animals which subsequently underwent the three different ablations. P values for the Primary Parietal vs. the Primary Temporal, Primary Parietal vs. Primary Frontal and Primary Frontal vs. Primary Temporal were .083, .125 and »-331 respectively. Using the same statistical procedure on the post-operative scores, these three groups yielded P (Probability) values of .190, .393 and .189 fo** the comparisons of Primary Parietal vs. Primary Temporal, Primary Parietal vs. Primary Frontal and Primary Frontal vs. Primary Temporal, respectively. Once again there are no significant differences in the error scores of the three groups of cats. Investigating the effects of the additional frontal and temporal lesions, using the U-test, yielded a P value of .192 for the difference in error scores between the animals who received the Secondary Frontal lesions and those which received the Secondary Temporal lesions. This is interpreted as indicating no significant difference in error scores attributable to the particular type of secondary ablation involved. To complete the analysis of the brightness discrimination data, the post operative scores of the six groups of cats were compared in the following combinations by means of the U-test: 24 1) Operative control ve 2) Operative control vs 3) Operative control ve 4) Primary parietal vs. 5) Primary parietal vs. 6) Primary frontal vs. 7) Secondary frontal vs primary temporal primary frontal primary parietal primary temporal primary frontal primary temporal secondary temporal These comparisons are based on the following percentage error scores: Gat No. Pre-Op. Post-Op. Operative control X L I I 20.6 2.6 XL 0.0 5.0 Primary temporal X L I I I 1.5 12.8 XL 4.2 21.8 LV. 9.4 18.7 LVI 4.0 4.0J XXVI 27.0 20.8 XXIV 2.9 51.5 Primary frontal XXXVI 5.1 20.8 X L I I 9.1 60.8 XLVII 1.7 5.0 XXXIX 18.8 25.0 XLVI 8.5 22.2 Primary parietal LIV 12.6 22.8 L V I I I 18.0 22.6 XLIX 12.0 42.0 Secondary frontal XXVI 52.8 XXXIV 45.6 LVI 24.0 Secondary temporal XXXIX 40.8 XXXVI 0.0 XLVI 7.5 XLII: 65.8 XLVII 0.0 LIV 27.5 XLIX 28.0 The seven comparisons outlined above yielded P values of .047, .042, .042, .190, .292, .189, .192 respectively. The only instances of significant differences in error scores occur when the operative control animals are compared to the Primaries Frontal, Temporal and Parietal. 55 In these cases the P.'values are .045, .047 and .042 respectively. This finding is in keeping with the previously noted observation that the actual ablation procedures resulted in an increase of error score. B. Behavioral Observations. In-; spite of the inability to differentiate the various cortical association areas in terms of their relation to brightness discrimina-tion, as tested in this experiment, there were noticeable differences between the three operated groups of cats. 1. Frontal Group. The most striking difference between this group and the other two was the amount of total activity. The cats with frontal ablations were hyperactive. This state was extreme in the immediate post-operative period but usually settled down to a moderate degree of hyperactivity. The frontal cats in this early post-operative period were reasonably alert yet gave the appearance of being confused. For the most part, there was some sensory and motor defect which was unavoidable with the surgical approach used for this ablation. This surgical approach of reaching the frontal poles from over the sensory-motor cortex was necessitated by the desire to leave the large frontal air sinuses intact to prevent possible infection. The amount of sensory-motor difficulty is reflected by the placing and hopping responses pres-ent (Bard, 1957-38). A l l animals walked normally once they got going but in starting or turning, one limb or more would slip out or the cat would stand on the back of one fore-foot. Cats with medial frontal ab-lations had a characteristic crouch in their hind legs, when walking they gave the appearance of waddling. 56i A l l frontal cate ate well initially yet once again the surgical approach over the sensory-motor area caused some defects. These were not immediately apparent as the animals were fed milk with pablum in i t which required only that the cat lap the milk. In two cases the cat reversed the curl of its tongue, being curled upward instead of down, yet was able to lap the milk. Later when the cats were given the regular mash, the difficulty in eating became apparent. There was an inability to co-ordinate tongue and jaw movements to get the mash into their mouth. In eating the daily ration, the cat could press its face into the mash and get enough to the back of the mouth where the swallowing reflex would take over and assist the eating, but when faced with the rewards which were small balls of mash approximately three-fourths of an inch in diameter, the cat had difficulty, often flattening them to the dish and thus being unable to get the reward. There was an element of sham rage in.one of the frontal cats (XXXIX) which supports earlier works (Kennard, 1945; Bard and Mount-castle, 1948), although two other cats (XXXIV and XLVII) demonstrated sham rage for a short period after frontal lesions. One other cat with only the orbital surfaces removed, did not manifest rage reactions, a l -though i t became extremely hyperactive, but showed obstinate progression described by Bailey and Davis (1942). These observations of sham rage and obstinate progression seem paradoxical as the cat (XXXIX) in which the head of the caudate was damaged did not produce obstinate progression as found by Mettler (1942), i t gave sham rage as found by Speakman and Babkin (I95O). And the orbital surface ablation in XLVIII gave obstinate progression but 57 no sham rage as stated by Kennard (19^5)• However, the significance of these observations on the two cats must remain tentative until a histo-logical study is completed on the brains. The lack of sham rage in the frontal cats in the maze may be attributable to the scarcity of stimuli, maze or extra-maze. Yet the one cat that had this symptom continued to demonstrate i t outside the maze until i t was sacrificed five and one half months after the frontal ablation. A frontal symptom, 'placidity 1, is perhaps related to emotion, yet i t is here being used to refer more to the plasticity of the cat than to reaction to the environment. The frontal cats could be picked up and held by the back of the neck for long periods of time without struggling. They would remain inert with a l l four limbs extended. Barria (1957) obtained frontal cats that were so placid as to justify the term 'cataleptic 1. Such a term could scarcely be applied to the frontal cats of this series. Frontally ablated cats were distinguished in the maze by their lack of emotion. Most cats and even the frontal cats before operation would show emotion by vocalizing, scratching at the alley walls or even the gates. Regardless of the number of successive errors and consequent lack of reward these frontal animalB continued to attempt each t r i a l . The animals in this group would walk around the surgery floor but would not manifest interest in any one object long enough to investigate i t . They walked up to the legs of tables or chairs and would move on to another and another in a restless fashion. This be-havior could be interrupted at any time by the experimenter moving about the room, then the cat would follow him until he stopped.. This 39 'following1 is another frontal characteristic and appeared in four cats (XLII, XLVI, XLVII, XLVIII). In the maze these cats were dis-tractable. This was defined in terms of the number of times the cat tried to re-enter the feed box once i t had returned varying distances back toward the starting box. 'Forced circling' as described by Kennard and Ectors (1938) was found in two cats (XLVIII and XXXIV). These investigators found unilateral destruction of area 8 of the monkey was necessary to eli c i t this behavior. All ablations here were bilaterally simultaneous. How-ever, inadvertent inequality of frontal ablation in a l l probability led to this circling behavior in these two cats. One last characteristic of this group was a slight de-crease in variability at the choice point, that is, there was a tendency toward stereotypy of choice following loss of the frontal lobes. 2. Parietal Group. With the exception of a slight paresis in the hind legs and the loss of hopping responses in cat LVIII, the animals with bilateral, parietal ablations showed none of the behavioral changes of the other two groups. The most noticeable characteristic of this group was the speed with which they recovered from the loss of nervous tissue. They were alert, normally active and hungry the same day on which the surgery was performed in sharp contrast to cats following frontal or temporal ablations. 3. Temporal Group. The total post-operative activity of this group was much less than that of the frontal or even the normal cats. In the immediate post-29 operative period these animals were extremely inactive and somnolent. They invariably curled up on a l l fours with their heads down and refused to move. When placed on their feet, they resumed this curled-up pos-ture immediately. Within two or three days the cats with the temporal ablations developed an apparent curiosity. This activity was more of a cautiouB investigation in which they walked about as i f they were 'stalking' something. A part of this type of behavior can be termed 'nosing' which is, perhaps, what Bucy and Kluver (19^0) refer to as 'smelling' or 'olfactory examination' in their investigation of the effects of a bilateral, temporal lobectomy in", the rhesus monkey. The temporally ablated cats in the immediate post-operative period did not appear to recognize the milk as food and many had to be force-fed. One often got the impression that the cats were actually blind as they would progress right up to an object before they would stop. It was not possible to demonstrate scotomata as these animals would not respond to moving objects for some time post-operatively. In fact, i t seemed the cats reacted in the opposite way to Kluver and Bucy'a (1929) hypermetamorphoais of the monkey. Pour of the cats (XXXIV, XLIX, LIV, XXXIX) sat and stared a great deal after ablation of their temporal cortices, while four of them (LIV, XXXIX, XLII, XLIX) had demonstrable transient hemianopias. These hemianopias were not apparent until i t became possible to attract the cat's attention with moving objects. Although there were the visual defects from temporal ablationB, these extensive lesions had no demon-strable effect on the cats' auditory ability. They responded normally to a l l sorts of auditory stimuli. 4p There was no noticeable paresis in the cats with temporal ablations with regard to eating. There was no doubt the cat could see the milk when confronted with i t , yet the animal paid no attention to i t . However, when the cat was force-fed milk by means of a rubber syringe, i t offered no resistance, in fact, acted as i f i t were hungry. Moreover, when faced with the regular mash later on the cat would eat, although i t gave the dish a good 'nosing1 over before i t attempted the food. In the cats of this group, the pupils were extremely dilated. There was, however, a good pupillary constrictor response to a bright light. In the immediate post-operative period two of these cats would seek out any dark portion of the room and curl up. They were often found in their living cages facing the dark wall and many attempts to get them to remain facing the lighted portion of the room failed. This symptom was transient, seldom lasting more than a few days, yet the pupillary dilation remained for a considerably longer period of time. Circling behavior similar to that found in the frontal cats appeared in five cats (XXXIV, XXXVI, XXXIX, XLIX, XLII). This circling was probably related to the hemianopias following their temporal abla-tions. A l l cats were able to compensate quickly for this defect. 4. Additional Ablations. The results here were contradictory. In most instances the additional l e s i o n B made a slightly greater deficit but this was only temporary and tended to disappear more quickly than the primary lesion symptoms of the same area. Cats with frontal lesions added to temporal, certainly showed the frontal symptoms of the same area. Cats with frontal lesions added to temporal, certainly showed the frontal syndrome; motor and sensory deficit, restlessness in the maze, stereotypy and following. These symptoms tended to persist. The secondary temporal lesions had an equally marked influence in depressing the in i t i a l frontal effects, yet the added deficits lasted a much shorter time with the frontal syndrome returning. 5. Additional Training. Incidental mention is made here of the effect of training on the results of subsequent cortical ablations. Two cats were carried on in the trials one month past the time they had reached the statistical criterion. In both instances there was much less deficit in regard to brightness discrimination following the second and subsequent operation. 42 DISCUSSION. Perhaps no concept is more firmly rooted in. neurophysiology than, the functional differentiation of various parts of the cerebral cortex. The past century of intensive anatomical research has settled beyond question the histological diversity of the cortical fields and of their connections with subcortical nuclei. These finds are amply borne out by a wealth of physiological and clinical evidence to support special functions of the majority of cortical areas. No one today seriously believes that the different parts of the cerebral cortex a l l have the same function. In this test situation no significant differences appeared as the result of focal, cortical ablations. However, behavioral differences, which were noted here, have been seen by other workers and i t is, perhaps, only a matter of time until experimental situations are de-vised with which these observations can be objectively quantified. A very good start was made by Jacobsen (1936), and has since been.con-firmed and elaborated by Harlow, Meyer and Settlage (1951). A. Frontal Group. From the brightness discrimination data this group appeared equally as disorganized as the temporal and parietal groups. The rest-lessness in the starting box of the frontal cats usually resulted in a rush to get to a feed box so that they leapt down whatever alley they were facing as the maze gate went up. This seems to indicate a genuine loss of ability, not merely .a loss of performance due to some motor or 43 sensory defect caused by the operation. It is true these cats became hyperactive but the speed with which they left the choice point was no greater than that of the speediest cat in any of the groups, and there-fore, they must have had sufficient time to make a discrimination. The failure of the frontal cats then could be as Wade (1952) suggests, either "loss of interest in the situation" or "hyperactivity" or a com-bination of the two in thiB study depending upon the extent of the abla-tion. This hyperactivity and distractability was shown by Wade (1947) to be the cause of the monkeys' inability to perform the delayed res-ponse rather than the former hypothesis of loss of immediate memory or retroactive inhibition. Such an explanation of activity as the cause of the defect in this experiment is quite a reasonable one. An adequate measure of the hyperactivity, however, cannot be de-termined in such a maze as was used in this study because of the paucity of extra-maze cues to which the cat is able to respond. It is, there-fore, not possible to observe the extent of the frontally lobectomized cats' 'over-reactivity' in the maze situation to the same stimuli to which the temporal cats pay no attention. But due to the hyperactivity of the cats and the lack of external cues i t is assumed the cats were responding to interoceptive stimulation. However, the number of times the frontal cats tried to re-enter the feed box once they had left i t , compared to the virtual lack of this behavior in the temporal cats, gave some indication of over-reactivity or distractability of frontal cats. This attempted re-entry of the feed box appeared to be an 'after-thought' or a response to some stimulus or other as the cat would be about to leave the alley when i t would suddenly turn and go 44 back. A characteristic noted in cats that had primary lesions made on the medial frontal cortex was the tendency to nearly reach the statis-tical criterion set for learning the discrimination, and then to revert to random responses to the light and dark stimuli. This behavior is quite different from the learning pattern observed in other cats of this series. Although checked in a preliminary analysis, the variability of errors did not show the steady decrement pattern described in the con-tinuity theory of Spence (1942) but rather tended to uphold the non-continuity theory of Krechevsky (1932). The reduction of variability in choice between right and left alley for this group is similar to findings of Morgan and Wood (1943) for cortically ablated animals. These workers found a decrease in variability of choice for rats with frontal and parietal lesions and and increase in variability for occipital rats. However, the reduction here was slight with only a small tendency to stereotypy in the frontal group. B. Parietal Group. Prom the lack of observable changes, i t would appear that the parietal cortex, which is assumed to be a nonspecific area since no studies have revealed in the cat any neuronal connections with this area other than a possible relation to the sensory leg area (Garol, 1942b), has none of the incapacitating features of frontal and temporal cortices. However, i t would appear to be a reasonable assumption that this area is involved as is the temporal area in the alteration of dis-criminative ability for the cat because of its close anatomical rela-45 tionships with the primary visual areas. However, i t has been shown in the monkey by Chow, Blum and Blum (1951) and Harlow, Davis, Settlage and Meyer (1952) that this area is not involved in loss of visual dis-crimination. C. Temporal Group. The discriminative changes brought about by the temporal lobe ablations in monkeys as seen by Kluver and Bucy were not quantifiable by means of the brightness discrimination test in cats used in this experiment. However, the subjective observations indicate a difference in behavior of the three groups of cats when they are confronted with the usual multiple stimuli of a conventional environment. There were two cats in this group (XLIX and LIV) and two in a preliminary study of twenty-two cats, which, most clearly resembled the temporal syndrome described by Bucy and Kluver. Furthermore, this syndrome persisted in these cats. They were slowed, had a glassy stare, nosed about in a cautious, stalking manner, and appeared disoriented in a usual situa-tion, except the maze. Contrary to the findings of Kluver and Bucy, i t was found in this experiment that the cats' "tameness0 was variable as was their emotional reactions. No changes were noted in diet or sexual behavior because the animals were individually caged and on a standard diet. It would be reasonable to conclude that the total stimuli of a given discriminate situation lead to these characteristics of the temporal lobe ablation, yet the stimuli in one dimension as used in this test are readily differentiated and not determinants of the res-ponses made in an uncontrolled situation. As with a l l operated groups there was no amnesia of the routine 46 required of leaving the starting box and progressing down one alley or the other until they reach the feed box, then returning to the start-ing box. It was, perhaps, due to their slowness that the temporal cats appeared more disoriented. They nosed about more and thereby usually saw both alleyways as they left the choice point, and i t was, perhaps, from this activity that they sooner realized their errors. The term 'realized' is inferred from the cats' hesitation at the choice point or by their stopping as soon as they entered the wrong alley and turning into the correct one. From this i t appears that somehow or other the cats were making a discrimination. D. Additional AblationB. The concept of mass action (Lashley, 1921) wa& not upheld here from the brightness discrimination results. In theory the degree of deficit increases with the more critical tissue that is removed. This is hardly a fair comparison of the results in this experiment with the theory of mass action as there was training and relearning going on between the Beriatim ablations of bilateral temporal and bilateral frontal cortices. Insofar as the brightness discrimination results are concerned, i t may be as Gthiselli (1938) suggested that such a simple discriminative ability as intensity differentiation may be mediated through the superior colliculi and not-be related in any way with cortical areas. From the work of Starzl and Magoun (1951) and Starzl and Whitlock (1952) which demonstrated that the frontal areas have a much larger projection from the reticular activating system than the other two association areas, i t is possible the loss of control from these frontal areas has a greater effect on the behavior of the 47; eat which is presumably operating at a lower and more habitual level. 48 SUMMARY Eighteen cats were instrumentally conditioned in: a Y.-maze, which was housed in a darkened room, to respond, either positively or nega-tively, to a stimulus light. After the cats had mastered the visual; discrimination, fourteen had simultaneous bilateral ablations of the various associative cortices, frontal, temporal or parietal. They were given, post-operatively, daily, training trials until they met the statistical criterion. Then the cats were subjected to secondary cortical ablations, either temporal or frontal, and once more retrained in the maze to respond to the appropriate light cue. The pre- and post-operative percentage error scores were used in the statistical analyses to determine the effect of these lesions on brightness dis-crimination. Daily subjective observations were made on the behavior of the animals and the changes were noted. The results from the sub-jective and objective observations made on the cats before and after cortical ablations appear to warrant the following conclusions: 1. There was no statistically demonstrable difference between the effect of frontal, temporal or parietal lesions on brightness discrimi-nation. Nor did additional secondary frontal or temporal lesions pro-duce significant changes. 2. There was a difference between pre- and post-operative cats with regard to brightness differentiation. J. There was a tendency toward hypomotility in the temporally lobectomized animals while in the frontal cats there was general hyper-49 activity. 4. The activity of the temporal cat ie what has been termed 'stalking' and the cats appeared to be cautiously curious about their surroundings. They nosed or 'smelled' objects a great deal. The frontal cats with their restlessness did not manifest interest in any one object long enough to investigate i t . 5. The frontal cats would follow any moving object, but the temp-oral cats could not be distracted from their measured, determined in-vestigation of the object at hand. 6. The temporal cats had widely dilated pupils but the frontal and parietal cats showed no alteration. Some of the temporal cats seemed transiently photophobic. 7. The frontal cats were much more dietractable. This did not show up in the maze due to the paucity of cueB but there was a marked difference outside the maze situation. 8 . The cats with medial frontal ablations approached the criter-ion for learning the discrimination and, later, did less well. The cats with frontal pole ablations, however, with their restlessness continued to make errorB at a fairly constant rate. 9. Emotionality changed in the frontal group. These cats became less reactive to successive errors. There was no observable change in emotionality of the temporal and parietal groups. 1 0 . The addition of temporal to primary frontal lesions only trans-iently altered the frontal symptom of hyperactivity. The secondary frontal lesion produced the frontal syndrome of more activity and this tended to remain. 50 11. According to the visual discriminations test, both the temporal and frontal groups seemed more disoriented in the maze. This impres-sion may be due to the cautious nosing of their surroundings or the purposeless activity of the frontal group. 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The projection of the frontal lobe on hypothalamus. J. Neurophysiol.. 1947, 10, 509-514. Woolsey, O.N. & Walzl, E.M. Topical projection of nerve fibres from local regions of the cochlea to the cerebral cortex of the cat. Bull. Johns Hopkins Hosp.. 1942, 71, #6, 515-344. Yerkes, R.M. & Watson, J.B. Methods of studying vision in animals. Behavior Monogr., 1., #2, New York: Holt, 1911, pp90. A P P E N D I X A STATISTICAL TABLES. Table I Trial Order to Avoid Scores due to Position Habits, Simple Alternation and Double Alternation. (Modified from Gellerman, 1925)• Table II Criteria for Determining Length of Bun Required for Statistical Significance with Different Numbers of Total Trials. (After Grant, 1947). Table III Discrimination Test Results. Primary Parietal: a, b, c. Primary Temporal: d, e, f, g, h, i . Primary Frontal: j , k, 1, m, q. Operated on then dropped: n, o, p, r. Table 17 Mann-Whitney U Test for n = 5, 4, 5, 6 andj. Table 7/ Critical Values for r for Sign Test. (From Dixon and Maasey, 1921)* Table I Trial Order to Avoid Scores due to Position Habits, Simple Alternation and Double Alternation. (Modified from GeHerman, 1935)' 1 R : . R R L L 2 L L R L L 5 L R R R L 4 L R L L R 5 R R R L L . 6 R L L R L 7 L R R L R 8 R L L L R 9 R R L R L 10 R R L L L 11 L R R L R 12 L L L R R 1? R R L R L 14 L R R L L 15 L R R L L 16 R R L L R 17 R R L R L 18 L L R R L 19 L R R L L 20 R L L R R 21 R R L L R 22 R L . R L L 25 L R R L L 24 L R R L R 25 R R L L R 26 R L L R L 27 L R R L L 28 L R L R R 29 R R L L R . 50 L R R L L 51 L R L R R 52 L L L R R 55 R R L L R 54 L L R R L 55 L R L L R 56 R R L L R 57 R R L L L 58 R R L R L 59 L R L L R 40 R L L R R 41 R R L L L 42 R L R R L 45 L R L L R 44 L L R R R 45 R L R R L 46 R R L L L 47 L L R R R 48 L R L L R 49 R L R R L 50 L R R L L 51 L L R R R 52 L L R L R 55 R L R R L . 54 L L R R L 55 L L R R L 56 R R L L R 57 R L R L L 58 R R R L L 59 L L R R L 60 L R L L R 61 R L L R R 62 R L R L L 65 L L R R L 64 L R R L R 65 R L L R R 66 R L L R L 67 IJ L R R L 68 I £ R L R R 69 R L L . R R 70 L R R L L 71 L L R L R 72 R R L L R 75 R L L R R 74 L L R R L 75 L L R L R 76 R L L R R 77 R L L R L 78 R R R L L . 79 L L R L R 80 L L R R R 81 R L L R L 82 L - R R R L 85 L L L R R 84 L R R L R 85 R L L L R 86 R L R R L 87 L I L L . R R 88 L R L J R R Table II Criteria for Determining Length of Run Required for Statistical Significance with Different Numbers of Total Trials. (After Grant, 1947). S Largest Number of Total Trials for which (Length of Run of Probability of a Run of at leaet S Successes) Successes is Below 0.05) P = 1/2 P = 1/ 5 P - 1/4 p s l / 5 2 2 5 5 9 4 8 19 42 5 6 21 75 205 6 10 60 284 1006 7 17 175 1125 5014 8 51 510 4486 9 59 1521 10 112 4551 11 218 12 429 15 850 14 I692 15 5574 Table III. In: the following table of results, R and L are right and left choices as determined by Gellerman'B table. The symbols z and / indicate whether the choice waa incor-rect or correct, and + and — mean", respectively, the animal was forced by leaving the gate to the incorrect alley down or the food was placed in that alley not according to random ohoice. Neither of these was considered a t r i a l . Cat Temporal Lobe Series XLIX : To be trained to get reward in dark alley. Date Trials 3 Date Trials _ J May 28 1953 R / 1* / L X L X R X June 26 29 R / R / L / R x L \ 27 30 R / R , L X L x L 29 June 1 L / R . R / L x R 1 30 2 L / L / L X R 1, R X July 1 3 R / R x L i R / L X 2 4 L / R x R L x L / 3 5 L / R / R L / L / 4 6 R / R/ L / L / R X 6 8 -5 R X R x L • R x R X 7 9 - , +2,3 R X R x R / R x R X 8 10 +3,4 R X R x R 1 R / R X 9 11 - +2,3,4 R X R / R / R / R X 10 12 +1,2,3,4 R / R / R 1 R/ R X 11 -5 / R / 1 R / 13 13 +1,2,3,4 R R R X 16 -5 R / / 17 15 -1,2,3 +4 R X R / R X R 18 5 20 16 +1,3 -2,4 R / R / R / R / R / 21 5 / / 22 17 -5 R R / L X L x L 23 18 -4 R / L x L X L < L / 24 19 L X R x R X L / L X 25 20 -4,5 L X R / L X L x L / 27 22 -1,2 L / R / L / L x R / 28 23 L / R / R X L x L / 29 24 L / R / L / R / R / 30 25 L / L x L / R / R / 31 -1,2,4 +3 -2 +3 +5 R L L R R R L R R R L L R R I X R / R R L R L L / x X \ X • I L/, L / X J L / L R L L L R L R L R L ! ! Operation Bilateral Parietal L L R R R L R R R L L R L / R L R R R L R L R L R L L L R R R R X .1 R / R L R R R L L L R R R R R L L L R L R L R L -R x R / / x x i Cat Temporal Lobe Series XLIX (Contd.) Date Trials _ J Date Trials 3 A u g , -1 1953 3 4 5 7 8 -h 10 11 -3 12 -2,3 13 Ik 15 -2,3 17 +3, -2 18 19 20 21 -2,3,5 22 2k 25 -1,4,5 26 -2,3,4 27 -2,3 28 L / R / R / L / L / R / L / L / R / R / Rx L / L x R/ L / Operation Bilateral Temporal I x x ', X X X * X •l X X X / L R L R R L L R R R L L R R L / X / X X X X \ X X / R x R x R / , •L / Sacrificed: Hasn't L R R R R R R R R R R R R R L R R L L / ) x x i x x ', I X R R L L R R L R R L L R R R R R R R R R L R R R L R R R L R R R L R L R L R met criterion. Cat Temporal Lobe Series LIV : Trained to get reward in light alley. Date Trials 3 Date Trials _ J June 15 16 17 18 19 20 22 23 2k 25 26 27 29 30 3 k 6 7 8 9 10 11 13 lU 15 16 17 18 20 21 July i x x I "I X X A R L R R L R R R •I L !l R / R x R x L L R L L R L L L R L L L L R L R L L R R L L / L R R L L R R L R R R L R Operation Bilateral Parietal \ x •I L R L L L R L < L i i ! x R / L R L R L R R L R L R L R L R 'I X X / X ! R L L R R L L R R L L R R L July 27 L / L /, R / R / L 28 L x R / L / L X R 29 R < L , L / R / R 30 R / L / R X L / L 31 L x L - R R - L Aug. 1 L x R - R - L - R 3 R x L - L - R - R k R / L x L - R - L 5 +3A L x L x L / L X L 6 L / R / L X R / R 7 R / L 1, L / R / R 8 L X R / R / L / L 10 L x L < R X L / R 11 R / R / L / L / R 12 R / k / L / R / R 13 L / L / R / R X L 22 Sacrificed. I X / X I Operation Bilateral Temporal Cat Temporal Lobe Series LVIII : Trained to get reward in light alley. Date Trials 2 Date Trials _ J July 16 1953 Aug. 17 18 20 21 22 23 2k 25 27 28 29 30 31 1 3 k 5 6 7 10 11 12 13 Ik 15 17 18 19 -3 -5 -3 +4,5 -3 / x X X X i X ; X X \ X R/ L x X / X i X / X X X X \ L L R L R L R L R L L L R L L L R L / R x R / L R L R R/ R x R L R L R L L R L I L R L R L R L L L R R R L R L R x X / X ', X / X X X Y> I X X / R / , R/, R / X / X •l X X ', X X i Operation Bilateral Parietal L R R L L R R R L x x ! x x L x R / i x / / X ! x / L L R R L R R R L i Aug. 20 21 25 L / L / R / L / Sacrificed. L / H / If Cat Temporal Lobe Series XXVI : Trained to get revard in light alley. Trials Date 1 > 3 I t 5 Date Oct. 10 1952 R / L / R / R X L / Nov. 2k 11 R x L / R / L , L x 25 13 *l R X L x L / R x 26 Ik L . R X R / L x R - 27 15 L , R X L x R / L x 28 16 R / L - L - R - R - 29 17 L x R X L - L - R - Dec. 1 18 L x L - L - R - R - 2 20 L x R - R / L X R - 3 21 L x L - R - R - L - k 22 L / R / R / L X L - 5 23 R / L X R / L / L - 6 2k L . R /- L x R X L x 8 25 i» / L / R x L X R / 9 27 R / L / X L L R X R x 10 28 L 7 R R / L X L x 11 29 R x R i; L x R ', L /, 12 30 L / L R x R 13 31 R x L / R X R X L / 15 Nov. 1 R x R 1 L 1, R \ L x 16 3 R x L X R / R L x 17 k L x L X R / L 1 . R x 18 13 R / L X R / R 1 L / 19 Ik L x R / L X R L x 20 15 L f. R / L 1, R 1 R x 22 17 L / R / R / .L X R / 26 18 1 R / L X L x R / L x 27 19 L x L X R / L / R / 29 20 R / L X L , R / L x 30 21 W R X L / R / L / 31 Trials — 2 R R L R R L R L L L R L R L R R R R R L R L R R / x X X X X / R / L x L L L R / L x I x i X X / R / L x L L R L L L / L L R L R L R / x X / X X / X R / L x / X / X X X I X 1 L R R L R L L L L L L L L L L L L L L L R L R / L x Operation Bilateral Temporal R / R x L / R x R / R x L / R x L x L / L ? L / R x L / R x L -R x L / Cat Temporal Lobe Series XXVI (Contd.) Bate , 1 2 Trials 3 1 5 Date 1 < 3 Trials 3 4 5 Jan. 2 1953 R / R / L / L / L < Feb. 9 R/ R / L / R / L / 3 R x L / R x R X L , 10 Operation Bilateral Frontal 5 R / L x R / R / L / 1U L x R / R x L / L / 6 R / R / L / L / X L / 16 L x R / R - L - L -7 R / L x R / R L , 17 R / R / L x L L R x 8 L , R x R / L ', 1» / 18 R / R / L x R / L x 9 R/ L < R X R L , 19 L x L X R / R x L / 10 L x L / R X R X L , 20 L x R X R / L x L x 12 R x 1' , R X L / X L , 21 R / L X L x R x R / 13 R x R / R / L W 23 R x R / X L x L x R / 14 R / l* / L / R ', R x 2k R / L R / L x L x 15 R / L / R X L L / 25 L x R i R / L / L x 16 R x L / R X R / 26 L / R R / L x R / 17 R / L / L / R X L , 27 R x R 1 L x L x R x 19 R / L , - L / R X R , 28 R / L X L x R/ L x 20 L x R / R / L / L / Mar. 2 L x R / R / L x L x 21 R/ L x L / R / R x • .- 3 - L x L X L x L / L x 22 L x L x R X R X W k - L x L X L x L x L x 23 R/ W L / R X 5 - L / L X L x L x L / 24 R x R x R X L / L / 6 - L x L X L x L x L x 26 R x L / L / R X / 7 - 2 , +1,3, L / L X W L / L / 27. l R x R X R / L / 4,5 -1» / L < 28 R / L x L / L / R x 9 -, +4,5 L x L X L x 29 R / L / R X R / L < 10 - L x L / 1* / L / L i 30 R / R x R / L / 1» / 12 - L x L X !• / L x L < 31 L x L > ! R / L / 13 - L x L X L < L x L < Feb. 2 L / R / R X R / L / 14 - L x L X L / 1» / L / 3 L / R / R / L / R < 16 R x R / L / R / L x k R / R / R / L / 1« / 17 L / R / L x L < R x 5 R / L / L / R X L < 18 R x L X L x R / R x 6 L ^ R / R / L - R / 19 R / R / L x L x L / 7 R / L / 1 L / L R/ 20 R / L / R / R x L X Cat Temporal Lobe Series XXVI (Contd.) Trials Date 1 2 3 4 5 Date Mar. 21 w R / L x L / R x May 1 23 L x L x R x R x R x 2 2k R x L x R x R / L x k 25 R x R / L x L x L / 5 26 L / L x R x R x R x 6 27 L / R/ L l L ^ R x 7 28 R x L / R / R / L / 8 30 L x R x R / L / L x 9 31 L x L x R x R x R:/ 11 April 1 L / L x R /, L /, R x 12 2 R x L x R /• R / L x lU 3 L / L / R / * , L / 15 k L X L x R x R / L x 16 6 R x R x L x L / R x 18 7 R / L 1, R / L x L / 19 8 R/ R / R / 1 / L x 20 9 L x L / R / R x L x 21 10 L x R x L x 1» / R / 22 11 R/ L / L x R x l» / 23 13 R / L x R / L L / 25 lU L x L x R / R / L x 26 15 L x R / R/ L x R x 27 17 R / L x L x R x R / 28 18 R / L x L x R / L / June 8 20 L / 1« / R / R / L x 21 L x R / L x R / R / 22 R / L x 1» / R / R x 23 L / R/ R / L x L L 2k L X L x R x L x R / 25 R / R / L x L / R x 30 R / L x W R x R / Trials _3_ -5 -4 -5 L X L X R / R / L L X L / R / X L X L R / L X L L / R R / L X L 1 R / L R / R / R L X L L X L X R 1 L X L L / L X L X L / L R / L X L / R X L L X R X R X R ', L L X L X L / R R L - R X R / L X R R / L X L / L / R R / L / R / R / L L / L / L / R X R L / R / L / R / R R / R / R / L X L L / L / R / L / L L / R / R / R / L L / R / L X L / R R / R / R / L / L R / L / L /. R / L L / R / R / L / R R / L / L / L / R Sacrificed. x X i x / X i x ! x \ I (0 H Cat Temporal Lobe Series LVI : Trained to get i t s reward in ligh t alley. T r i a l T r i a l Date 1 2 3 ^ 5 Date 1 2 3 4 5 Operation Bil a t e r a l Frontal L / L / R/ L / R / R/ R/ L X L / R / R x L / L / R X R X L / L / R/ R/ L X L / L / R/ L / R X R x L / L / R X R X L / L / L / R/ R X R/ R X R/ L / L / L / L / R/ L / R / L / L / R/ R/ R/ Sacrificed. June 19 1953 L-X R x R / h , L X Aug. 6 20 L / X R x L X R / R - 10 22 R R / ..L / L / R X 11 23 L I R x R X L X L / 12 24 L R / L X R - R - 13 July 6 R / R x L X L / L - 14 7 R / L - R - R - L - 15 8 L / R - L mm L - R - 17 -1,4,5 9 L / L - R - R - R •- 18 10 R / L - R - R - L - 19 11 R / R / L / L 1. L / 20 13 L / L , R / R / R X 23 14 L / R / L / L , R / 15 R / L x R / R / L / 16 L / R / R / L / L X 17 L / L / R / R x R -18 L / L , R / L - R -20 R / L / R X R - L -21 L / R / L / L - R 22 L / L , R / R / L / 23 R / R / L / L , R X 24 R / L , R / L / L / 25 R / R 1, R / L ^ L / 27 L / L / R / R/ L / 28 Operation Bila t e r a l Temporal 31 L / L 4 R / R / L / Aug. 1 L / R ^ R / R / 3 R / L / L / R / R X 4 R / L , L / R / L / 5 L / L / R / R / L / Cat Temporal Lobe Series LV : Trained to get its reward in dark alley. Date Trials 3 Date Trials __2 June 18 1953 + 19 +5 20 22 23 July 13 lit 15 16 17 18 20 21 22 23 • 2k 25 27 28 29 30 31 Aug. 1 3 k 6 7 8 10 R L L R L L L R L L L R L L R R L L R R L L x / X j L R R R/ R x L x L !, R / L L L R-/ L x x ! R L R L R L L L R L / X / X / X ', X I X I X I / X X ! X X X X X I L /t R / L L R -R/ X / Aug. L R R L L R R L R L R L L L R R L L R R / x / X X 1I X i X \ X I Operation Bilateral Temporal L x Rx L x R- R-R/ L / L / R- R-L / R X R/ L / L / L / L / R/ L x R/ 11 12 13 22 R / R / L / R / L / L / L x L / R / Sacrificed. L / R / R / R / R/ L / of & H H H Cat Temporal Lobe Series XLIII : To be trained to get reward in dark alley. Trials Date 1 2 3 4 Feb. 1 2 1953 L / L x L x R / R 13 R x R / L - R - L 14 L / R x R x L 4 R 16 L / R / R x L , L I T R x R x L 1. h , R 18 R x R / L / R < L 1 9 L /, L / R / R < L 20 L / R x R x R / L / L 2 1 L x L / R x R 23 R/ L x L / R 2k R x L / R x L x L 25 R / R / L / L 2 6 L / R / R x L x L 27 R x R / L 1. L / R 2 8 R x L < L R x R March 2 L < R / R/ L x L 3 L . R x L x R / R k R / R / L x L x R 5 L / R x R / L x L 6 L < R x L . R x R 7 L / L /, W R / R 9 R x R/ L x i« / R 1 0 L x L x R x R / L 1 2 L / R x L x L x R 13 R x R x L x L x R Ik R / R / L / L / L 1 6 R x R/ L / R / L 1 7 L / R X L x L X R 1 8 R x L / L / R x R 1 9 R x R / 1» / L / L 20 R / L / R / R x L Date Trials _ J x / X ! X ', X X / X X X ! X ', X / April March 21 23 24 25 2 6 27 2 8 30 3 1 1 2 3 k 6 7 8 9 10 11 13 14 15 16 17 18 20 21 22 23 24 25 30 L L 1 x / X X ! R L L R L L L R L L R R R L L R l / R L L R L R L / x L / R x R/ L X R x L L L L R L L / R^ Operation R/ 1 / .1 R L L L R R R R / R L < R 7, L < R < R / •I R L R / , x I X • I R < R / Control, 1 / R L R L R L R R R L L L R L R L R L •I R 7/ L / x / X / X / X / X X ! R < R < R i R < L < L // L // R / R R L L R L L L R R L L L R L L L R R L L R R L L R R L R B / Cat Temporal Lobe Series XLIII (Contd.) Trials Trials Bate 1 2 3 h 5 Date 1 2 3 May 1 L / L / H / R / L x 2 L / L / R / L / R / 4 R / L / L / R / R / 5 R X L / L / R / L / 6 R / R / R / L / L / 7 L / L / R / L / R / 9 R / L / L / R / L / 11 L / R / R / R / L / 12 L / L / L / R / R / 14 L / R / R / L / R / 15 R/ L / L / R/ R/ 16 R / L / R / R / L / 18 L / L / L / R / R / 19 L / R / L / R / R / 20 R / R / R / L / L / 21 L / L / R / L / L / 22 Operation B i l a t e r a l Temporal 26 R / L x L / R / L / 27 L / R x R / L / R x 28 R x L / L x L / R / 29 R / R / L x R / L x 30 R / R / L / L / L / June 1 L / R / R / L / R / 2 L / L / L / R / R / 3 R / R / L / R / L / 4 L x R x R / L / L / 5 L / R / R / L / L / 6 R / R / L / L / R / 8 R / R / L / R / L / 9 L / L / R / R / L / 10 Sacrificed Cat Temporal Lobe Series XL ; To be trained to get reward i n light alley. Date Trials Date Trials 2 Jan. 29 1953 R / L / R/ R / L X March ,3 30 R / R / R x L - L • a 5 31 L X L - R - L - L - .' 6 Feb. 2 L X R / R/ R X L X 7 3 L X R X L x L / R X 9 4 R X R / R 7 L X L X 10 5 R / L X L x R / L X 12 6 L X R / X R / L X L X 13 7 R / L L x L X L X 14 9 L X L / L x L X L X 16 10 L / L / L / L / L / 17 11 L / L / R x R X R / 18 12 L / L / k / R X R / 19 13 -5 R X R X L / R X L / 21 14 L / R / R / L / X L / 16 L / R X R / L L / 17 R / R X L x L / R / 18 R / R X l > : R / L / 19 . L / L / R / R / L / 20 L / R / R / L / L / 21 R / L / L ^ R / R / 23 R / R / L / L / R / 24 Operation Control 26 L / R . L / L / 27 R X R / L / L / R / 28 R / L . L / R / L / March 2 L / H / R / L / L / Operation Bilateral temporal. L X R X R - L - L L X R X L- R - R L X L / X L l R X R R X R L , L / R L X L / R / R / L L / R X L / L R R / R / L / L - R R / R / L , L / L R / R / L , R / L L X R / L / L / L R / L / L / R / R R / R / L / L / L Sacrificed. ! >-3 & H (D M M H !3* Cat Temporal Lobe Series XXXIV : Trained to get reward i n dark alley. Date Trials 2 Date Trials 3 Nov. 19 1952 R X L / R x R / L / Dec. 26 L / R 20 R X R / L x L X R X 27 R x R 21 L / L / R x R X R / 29 L / L 2k R X R X L / L / R X 30 R x R 25 L / R X R x R X L / 31 R x R 26 R X R X R x L / R X Jan. 2 1953 R x R 27 R X R X L / R X R X 3 R x L 28 R X R X R x R X L / 5 R x L 29 R X R X R x R / R X 6 R x R Dec. 1 R X R i R / L X R / 7 R x L 2 R / R L,x L X R X 8 L / R 3 R / R / L x L X L X 9 R x L k L X L / R / R ', L X 10 L / L 5 L X L / R / R L X 12 R x L 6 R / L / R / R L / 13 R x R 8 R / R / R / L / L / Ik R / L 9 L / L / R /, L 1 L / 15 R / R 10 R / R / R / L / L / 16 R x R 11 R X L / L . R / L / 17 R / R 12 R / R / R 1 L / 19 R / L 13 R / R / L / L 1 L / 20 L x R 15 Operation Bilateral Temporal 21 R !, L 17 R X R X W R X L , 22 L . L 18 L / R X R x L / L , 23 R / L 19 R X R X W R X L / 2k R / R 20 L / L / R x R X L / 26 R / L 22 R X R X L / L / R X 27 L / R 23 R X L / R x L / L / 28 R / L 2k R X R X L / L / R X 29 R/ L X X / X X X 'l X / X \ X X X I R L R L L L R R L R R R R R R L L L R L R L R L " j x I X / L / R/ R / L L R L R L R R L R L R R L L R R L R R L R R R L R x / X / X X / X / X X ', X ', X X / X X I / X I X ! X J R L L L L L R L Cat Temporal Lobe Series XXXIV (Contd.) Trials Date 1 • > 3 4 t Date Jan. 30 H / R / R/, L , Mar. 13 31 L / L / R / L , L 7 / 14 Feb. 2 L x R / R / R / L / 16 3 Operation Bi l a t e r a l Frontal 17 9 R x R X L , R - L - 18 10 R x R X L < L X 19 11 L / R X R x L / R X 20 12 L . L / L L R x R X 21 13 R , R / L / R x L i 23 14 L / R X R x L / L 24 16 L / R X R x L < L 1 X 25 17 R x R X L i L / R 26 18 R x R X L / R x L / 27 19 L < L / R x RX L / 28 20 - L / R X R x L / L / 30 21 R x L L 1. R x R X 31 23 R x R L / L < R X Apr. 1 2k R / L / R x !• / L / 2 25 L < R X R x L / 3 26 L / R X R X L / R X 4 27 R x R X L 7 L / R X 6 . 28 R < L / L / R x L ', 7 Mar. 2 L / R X R X L / L 8 3 L x R X L / R x R X 9 k R x R - L - L - R - 10 5 E fi R X R X L / L / 11 6 -k L / R X L / R / R 13 7 L / L / I* / R x R X 14 9 -1, - 2 , 3 , R x R / R / R / R / 15 4,5 18 10 +1 , 2 , 3 , 4 , 5 R / R / R / R / 12 +1, - 2 , 3 , 4 , 5 R / R X R x R / R / Trials _ J I x x J>, L / R R L L R L L L R L L R R R L L R R L L Sacrificed. L / L / R R / L - L - L R / L / R / L R / L / L / R L /, L / R X R R / L X L / L L /, R X R X L R / L / L / R L < R X R X R L < R X R / L R / L / L X L L / R X R / R R / L / L / R L / R / R / L R / R / L / L L < R / R X R L / R / L / R D / R / R / L L / R X R / L L / R / R / L R x L / L / R L / R X L / L R / R X L / X L L < R X R L R / L / L / R L /, L / R / R L < R / L / L L / R X R / L R/ R / L / R / ) X / X / X ! X ! ! Cat Temporal Lobe Series XXXIX : Trained to get reward i n dark alley. Date Trials __2 Date Trials — 2 Jan. 12 1953 R / L X R / L X L X Feb. 20 L / R / R / L / L 13 R / R R / L X L X 21 R X L / L / R / R 14 R X L L X R / R ', 23 R X R X L / L / R 15 R X L 1 R X L / L 2k R X L / R X L / L 16 R / L L / " R / R 1 25 L / R X R X L / L 17 R X L L / R X R X 26 L / R X R I L / R 19 R X L 1 L / R X R X 27 R X R X L L / R 20 L / R X R X L * L X 28 - R X R X R 1 R X R 21 R X R X R X R X R X March 2 -3,4,5 L / R / R X R / L 22 + R X R / R / R X R / 3 -3 L / R X R / R / L 23 R / R X R / R / R X k -3,4 R X R X R X R X R 24 R / R X R / R / R / 5 R X R X R X R X R 26 R / R / R / R / L X 6 I, +4 R X R X f R X R / R 27 R / R / L X L X R / 7 -, +1 R / R / R / R X R 28 R X L. / R X R / R / 9 -, +1>5 R / R / R X R X R 29 R / L X R / R / L / 10 +3,4, -5 L / L / R / R / R 30 R X R / R / L X L / 12 -4 L X R X L / R X R 31 L / L X R / L X L X 13 R X R / L / L / R Feb. 2 L / R / R / X R X L X lk -3 R X R X R / L / L 3 L / R / L L / R / 16 R / R X L X R / L 4 R / X R / R / L X L / 17 L / R \ L / L X R 5 R L / X L / X R X f L / 18 R X L L / R ', R 6 L / R R L / R / 19 R X R 1 L / L L 7 R X L / L / L / R / 20 R i L X R / R X L 9 R / R / L / R / L / 21 L R \ L / L / R 10 R / R / L / L / L / 23 L X L R / R X R 11 L / R / R / L / R / 24 R X L 1 R / R / L 12 Operation B i l a t e r a l Frontal 25 R ', R X L / L X L 16 L / R - R - L - L - 26 • * L / R / R ', R 17 R / R - L - L R - 27 • * / R / L / L R 18 R / R - L - R - L - 28 R 1 L / R / R 1 L X 'l X X X ', X X / X X \ X X ', X / i I Cat Temporal Lobe Series XXXIX (Contd.) Trials Trials Date Mar. 30 31 Apr. 1 7 8 9 -5 10 -3 11 -2 13 -14 -15 -17 -18 20 -21 -22 23 24 25 30 May 1 2 4 5 -5 6 7 . 8 -5 9 11 12 14 -4 +2,3,5 L1, R / R / L / ^1, L / L / R / R / R/ Operation Bilateral Temporal R / L / R / L X L < R x R x R x L / L / L / L / R x R X R / L / R x R / L / R x R x R / L , R X R x R / R x R < R X R x R X R x R x R X R x R x R x R / R X R / R x R / R x R X R x R x R / R / R X R / R./ R/ R x R / R x R / R x R x R / X R / R / L > I L / R R / L , R / R / L ', L x L , L / R x L R / R/ R x L / L X R / R x L x L x R X R / L x L / R x R / L / L / L x L / R / R x L / L ^ R / R / R x L . L , R X R x •R x R / R / L / L x L / L x R x L / R/ L < L x R>! R / R / L x L / R X l /, w R / R x R X l ^ L x L x L / R / R/ L / R x R x R X R x Date 1 2 3 I 1 5 May 15 -2 R / L x L / R x 16 -5 R x L / R x R X R / 18 Ll L x W R / R / 19 L /, R / L x R / R x 20 R / R / R / L X L x 21 L / L x R / L / W 22 L x R x R / R X L x 23 R / L x L / R x 25 R / R x R x L X L x 26 R/ L / R X L x 27 L x R x L X R / 28 R x L x L / L X R x 29 R / R / L / R / L / 30 R / R x L x L X L x June 1 L / R / R / L X R x 2 L X L x L x R / R / 3 R /- */, L x R / L x 4 L x R / L x L X L x 5 R x L / L x- R X R x 6 R / R / L x L / R / 8 -1 D / L x R X L / 9 -3 L x Ll L /, R / 1" / 10 L 1, R / R / L X L / 11 R / L / R X R x 12 -3 R x R x R / L / R / 13 -4,5 R x L / R x R X R / * 15 L / R x R / L X L x 16 M H < R / L / R / 17 -5 R / R / L x L X L /, 18 R / L x L / R / L / . 19 L / R / R x L X L x t H a H H H Cat Temporal Lobe Series XXXIX (Conta.) Date Trials _ 2 Date Trials _ ^ June 20 L / R / L / R X R / July 25 22 R X R / L / L X R / 27 23 L X R X R / L / L / 28 2k L / R X L / R / R / 29 25 L / L / L / R / R / 30 26 R / R X L / L / R / 31 27 L X L i R X R X L X Aug. k 29 L / R L 'r L i R i 30 R X R / L L R July : , i R X R / L 1 L 1 L / 2 R X R 1 L / R X L / 3 L / R / L / L ', R / R X L L / R R X 6 R X R / L / L 1 L 1 7 R / L 1 R X R X L 8 - U L / X R X L / R / R 1 9 L L / R / R X R X 10 R / L / R / R / L / 11 R X R / L / X L / L / 13 L / L / R R / R / Ik L X R / L / L X R / 15 R X L / R / R X L / 16 L / R / R / L / L X 18 L / L / R X L / R / 20 R / L / R / R / L / 21 L / L / R / X R / L / 22 L / L / R R / L / 23 R / R / L ', L X R / 2k R X L / R L / L X X X i j Sacrificed. R/ R^ L R L R L Cat Temporal Lobe Series XXXVI : Trained to get reward in light alley. Date Trials _ J Date Trials Dec. 8 1952 9 10 11 12 13 15 16 17 18 19 20 22 23 2k 26 27 29 . 30 31 Jan. 2 1953 3 5 6 9 10 11 13 Ik R L R R R R R L R L R R R R R L R L ] R/ L x / X X / R L R R 1 x ' j X i R R R L R R R L L R R R L L L L R L R L I X X / X / X A L / L L R L R L L L R R i x / X j L L L R R L R L R l / L L R L L L R L .1 R R L L L L L L L L L L R L R L R L R L L L L L L Operation Bilateral Frontal. L / R x R x R/ i x R / R x R x L ! , R x L 4 L / L x R/ Jan. 15 16 17 19 20 21 22 23 2k 26 27 28 29 30 31 2 Feb. 3 k 5 9 10 11 12 13 Ik R / R R L R L R R R L / x I X .1 R R L L L L X R L / L L / L L / L R X R L / L L X R L / L R / R L X R R / R L / L L / R R / R L / R R / R R / L R / R X \ X / X i L R R R L R R .1Operation Bilateral Temporal L h R < R < L < R // L i L / L x 1 x .1 L / R / R < L/, R / R i R < L < R / Sacrificed. L / R x L / R i R / L ! , R < R ^ L / Cat Temporal Lobe Series XLVII : To be trained to get reward in light alley. Date Trials __3 Date Trials 3 March 5 6 7 9 10 12 13 14 16 17 18 19 20 21 23 2k 25 26 27 28 30 31 April 1 2 3 k 6 7 8 1953 +4,5 +2,3 -1,2 -5 -5 -3,4,5 +3 -, +1,2 ', x x X X { X X X X / X X X X / X \ R x R x R x R/ R R R R L L R R R L L R L R L R L L L L L R L R R/ R/ R / R L L L L / x X X X X / X i R x R x / X X X I X L R R R R L L L R L L R R R R R L R L R L I L R R R R L L L L L R R R R R L L R R L L R R L L L R L L / x / X X X X X X / X X X i X X X ! April 9 10 11 13 14 15 17 18 20 21 22 23 24 25 30 1 2 4 5 6 7 8 9 11 12 14 15 16 18 May L L R R L L / R x L / i L i L / L 7, L !, M R / i Operation Bilateral Frontal. j R 4 L / L x L i R < R L L L L L R L L L R L R L L L R L L L L R L L .j R / L L •I R /, R < R L R L s/, R R R R R L L L R R L R R L L R R R R L L R R / x .! R R L R R L L R R L L R L i R / Cat Temporal Lobe, Series XLVII (Contd.) Trials Trials Date 1 2 3 ^ 5 Date 1 2 3 May 19 L / R / L / R / R / 20 Operation Bilateral Temporal. 23 L / R / L / L / R / 25 R / R / R x L / ' L / 26 R / L / L / R / L / 2? L / R / R / L / R / . 28 R / L / L / L x R / 29 R / R / L / R / L / 30 R / R x L / L x L / June 1 L / R / R / L / R / 2 L / L / L / R / R / 3 R / R / L / R / L X 4 L / R / R / L / L X 5 L / R / R / L / L / 6 R / R / L / L / R / 8 R / R / L / R / L / 9 L / L / R / R / L / I O L / R / R / L / L / u R / L / L / R / R / 12 R / R / L / L / R / 13 R / L / R / L / L / 15 L / R / R / L / L / 25 Sacrificed. Cat Temporal Lobe Series XLII : To be trained to get reward i n light alley. Date Trials Date Trials _2 May 20 1953 L / L / R x R / L x June 1 L / R X R / L x R 21 L x R x L x R / R / 2 L X L / L / R / R 22 -4,5 R / L x L x L X L X 3 R X R X L X R x L 23 L x R / R x L X L x 4 L / X R / R X L x L 24 L x L x R x L X R / 5 L R / R / L X L 25 R / R / L x L X R / 6 -5 R / R / L X L x L 30 -4,5 R/ L x L x L X L x 8 - L X L X L X L / L 1 - +3,5 L x L x L 1, L X L 1, 9 _ L / L X L X L x L 2 +2,3 -5 L x I* / L / L / 1" / 10 -2,3,5 L X L / L / W R 4 -3, +5 R x L / R x R X R / 11 R / L / L / R x R 5 -2,3 +4 R x R x R x R / R x 12 -5 R / R / L X L x L 6 +1,3 +4,5 R/ R x R / R / 13 R / L / R / L / L 7 -3,4,5 L X L x L / R / I» / 15 L / R / X R / X L x L 8 L x L l R x R X R / 16 L / R R L L R 9 R / L / L / R X L / 17 R X R ', L / L / R 11 L , R x R / R / L / 18 R / X L L / R x L 14 L , R / R / L / R / 19 L R / R / W L 15 R / L , L / L / R / 20 L / R / L / R x R 16 R / •L / R / R / L x 22 R / R / L / L / R 18 L x L , R / R < 23 L / R / X R / L / L 19 L 1, R / L /, R / R / 24 L / R L / R x R 20 R / R / R / L / L , 25 L / L / L / R x R 21 L , L ^ R / L / L , 26 R / R / L / L / R 22 L , R / R / R / L , 27 L / L / R / R x L 23 L / R x L / L / R / 29 L / R / L / L / , R 25 Operation Bi l a t e r a l Frontal 30 R / R / L / L , R 28 R x L / L / L / R x July 1 R / R / L / L / L 29 -3,5 +4 R x R x R x R / R x 2 R / R / L / R x L 30 R / R x L - L L - 3 L / R / L / L / R / X / X X X X / X ! X / X / X I Cat Temporal Lobe Series XLII (Contd.) Date Trial 3 Date Trial 3 July Aug. 4 1953 6 7 9 10 +2,3,4 11 -3 13 +2 14 -1,3,4 15 -2 +4,5 16 +1,3,4,5 17 +1, -2 18 -1,2 +3, 4 -4 20 21 22 23 24 25 -5 -4 2,4,5 4,5 27 -1,2 +3 28 -1,5 29 -2,3 30 +2,3 -4,5 31 -2,4 1 -5 3 -4,5 4 -4 +5 5 +3 R / L / L / R X R / R/ R/ L / L / L / Operation Bilateral Temporal R / R L R R R R / R x / x X X X X X / X X \ X R i R / R R R R R L R L R R R R L R R R R L L L R R R R R R R R x / X ',. X I X X / R x R x R -R -R x R / R x R / R x R L R R R R R L R R L L L L L R x R/ R -R -R ! , R 7/ R i R / / x / X X X X X X / X X X / X Aug. 6 -2 +3,4 L x L x L / L / L 7 +1,4 +2 L x L / L / L x L 3,5 t / / L / 8 +1 -2,3 L x L L 5 10 -4 L / L / R X R x R U -3 R x R x R L x R 12 R/ L /, L R/ R 13 L , L , R 1 R x L 14 L / L , R 1 L / R 15 R/ L , L / R x R 17 -5 R x L , L / R x R 18 R / R / R X L / L 19 L x L / R X L x R 20 L / L X R / R / R 21 R / L / L / R/ L 22 L / R x R / R - L 24 L / L / L / R / R 25 R / R / L / R 26 R / L x L / L X R 27 R / L x R / R x L 28 L / W L / R / R 28 Sacrificed: X / / / X Hasn't met criterion. Cat Temporal Lobe Series XLVI : To be trained to get reward in dark alley. Date Trials _ J Date Trials __2 Mar. Apr, May 24 1953 25 26 27 28 30 31 1 2 3 k 6 7 8 9 10 11 13 Ik 15 17 18 20 21 23 2k 25 30 1 2 k 5 -3,4 / x X X •I M L 7/ R / , L/, R // L / Operation / x X / X X X / X .! R L R L L L R L L L R / R / L / May 6 L x L / L / 7 R / R x R x 8 L x L / R x 9 R / R / L / 11 R / L / L / 12 R x R x R / 14 R / L / R / 15 R x R / L / 16 R / R / L / 18 R / R / L / 19 L / L / R / 20 R / L / L / 21 R / L x L x 22 R / R / L / 23 L x L / R / 25 L / R / R / 26 R / L / L / 27 R / R / L / 28 R / L / R / June 1 L / R / R / 2 L / R x L / 3 R x R / L / 4 Bilateral Frontal,. 5 R / L / L / 6 R / L / R x 8 L - L - R - 9 L / R x R x 25 . L / L x L / R x L / L / L / R / R x L / R x L / R / / R / L x L L R R L L R L L L R R L R x L R R R L R x I X I X .! R L R R R R L R L R L R L L R R L L R R R R L L x .j R 4 M L !i L 4 R / L L R L / x X \ X ! X Operation Bilateral Temporal L L R L L R R L R L R R R R L / L !l L 4 M R i L < L ft R / R R L L L R L L Sacrificed. H M Cat Temporal Lobe Series XXXIII : Trained to get reward in dark alley. Trials Date 1 2 3 4 5 Date Oct. 27 1952 R 7 L x R/ *!, L / Dec. 10 28 L X R / L x L / R x 01 29 R x L x R x L , R x 12 30 R x L 1, L / L 4 R x 13 31 L x L / R x L / R x 15 Nov. 1 R x L x L J, R x R / 16 3 R x R x L / L / R x 17 4 L 1. R x R x R x L x 18 13 L . L / R x R x L x 19 1 4 R / L X R / L x R x 20 15 L x L x R x L x R/ 22 17 R / R / L x R / L x 23 18 R x L x R / L X L x 2 4 19 R / R/ R / 26 20 L x R / L x L x R / 27 2 1 R / L x L x R / R / 29 2 4 L x L x R / L x L x 30 25 L x L / L x L x L / 31 26 L x L x L x L x L / Feb. 1 4 27 L / L / L / R x R x 28 L x L x L x L / L x 29 L', L x L', R x L / Dec. 1 L / R x L / L / R x 2 L /, L L R x R x R/ 3 L . L , R x R / R / 4 R / L /, R x L x L x 5 L / R / R / L / R / 6 L x R / R / L x L / 8 R / R x R x L / L < 9 L / L / R / L x L / Trials __2 R R R R R L R/ L/, R / / R  !, R/, R !, L/, R/, R / R 4 L ^ R ^ R ^ R ^ R/, L/, R ^ x J R L R R R L L/, L ^ L ^ R ^ L /: R x R ^ L R R L R L L L R L R L x J R L L L R L L L l x .1 L / / x x I X / X L // L // L / L x Operation Bilateral Temporal. Sacrificed. Cat Temporal Lobe Series XLVIII : To be trained to get reward in dark alley. Date Trials _ 2 ! L 5 Date 1 4 L x June 22 R x R L - 23 R R - 24 L / R L x 2k W L L / 26 R x R R x 27 L / L R - 29 L L / 30 R X R R / July 1 R / R R / 2 R Trials 3 May 21 ,1953 L / 22 L / 23 L x 25 R / 26 R / 2 7 - 4 L / 28 -3,4 R x 29 R x 30 -3 ,4 ,5 R x June 1 - 4 L / 2 -1,3 +2 R x 3 - 3 , +2 R x 4 - +2,3 R x 5 - R / 6 -3,4 R / 8 R / 9 -1,2,4 R / 10 - +3,4 R x 11 - +2,3,4 R x 12 - +2,3,4,5 L x 13 +1,2,3,4 -5 L / 15 -1,2 +3,4,5 L x 16 L -17 -1,4 -2,3,5 L / 18 - 3 , 1 * R x 19 L / 20 L / L x R / L / R / R / R / R / L x L -R / R / L / L / L / R x L / R x L / R x R x R x R x L / R x R x R x R / R/ R / R/ R / R/ R x R x R x R x R -R x R x R -R / R / R / R / R x R x R / L x. R / R / R / L x R x R / R / R / R / R / L / L / L / L / L / L / L x L -L x L x R x R x L x L x L / L / L / L - L ^ L / L x L / L / L / R/ L / L / Rx R/ L / L / Rx L / R X R/ Aug, 3 9 L x L -R x L -L x L / R / R / L / R / L x Operation Bilateral Orbital. Sacrificed. Cat Temporal Lobe Series LII : Trained to get its reward in light alley. Date Trials 3 Date Trials 2 July 11 1953 R / L X L / R / R / 12 R / R X L / L / R X 13 R / L / R X L X L X 15 L X R X R / L X L X 16 L X R X R / L X R / 17 R X R / L X L / R / 18 -4 R / L X L X L X L X 19 -2 +3,5 L X L X L / L X L / 20 -1,4,5 L X L / L / L X L X +2,3 / / / / 22 +1 -2,3 4 L L L L L X 23 -4,5 L / R X R X R / R X 24 -3 L / R X ' R / R / R X 25 L / L / L / R / R / 2 R / R / L X R / L X 3 -2 L X L / L / L / R I 6 R / R / L X L / L 7 R / L / R R / L / 8 L / R / L L / R 1 9 L / L / R 1 R / R X 10 R / L / X R / R / L / 11 R / R L / L / L / 13 L / L / R / R / R / 14 L - R - L - L — R 15 R / L / R / R / L / 16 L / R / R / L - L -18 L / L / R / L / R / 20 R / L / R / R / L / July 21 22 27 28 29 1 22 Aug. L / R / L / L / R / Operation Bilateral Temporal. L x L -R x L x Sacrificed. L X R - R - L -R - L - L - R -L - L - R - R -R - R - L - R -Cat Temporal Lobe Series LIX : Trained to get its reward in dark alley. Trials Date 1 - 2 3 July 23 1953 R / R/ L / R/ 2k R / L / R / L x L x 25 R / a / R x L / L / 27 L / L / R / R / L / 28 L x R / L / L x R / 29 R / L X L / R / R / 30 R / L X R / L / L / 31 L / L / R / R / L X Aug. 1 L x R x R / L / R / 3 R / L x L / R / R / U R / L x L / R / L x 5 L / L / R / R / L / 6 L x R / L / R / R x 7 R / L / L X R / R / 8 L / R / R x L x L / 10 L / L x R x L x R / 11 R / R X L / L / R / 12 R / L / L / R x R / 13 L / L X R / R / L / 14 L x L / R x L / R / 15 R / L x L / R / R x 17 R / L / L / R / L / 18 R X R / R / L / L / 19 L / L / R / L / R / 20 L / L / R / R / R / 21 Sacrificed. Table 17. Probability of Obtaining a U not Larger than that Tabulated in Comparing Samples of m and m (Tabulated P's are for a one-tail test of significance) n . 2 1 2 u. 0 .250 .100 .050 l .500 .200 .100 2 .750 ,4oo .200 2 .600 .550 4 .500 5 .650 n = 4 IK 1 2 2 4 U 0 .200 .067 .028 .014 1 .400 .122 .057 .029 2 .600 .267 .114 .057 2 .400 .200 .100 4 .600 .214 .171 5 .429 .242 6 .571 .242 7 .442 8 .557 T Table IV n = 5 m V 1 2 3 4 5 0 .167 .047 .018 .008 .004 1 .535 .095 .036 .016 .008 2 .500 .190 .071 .032 .016 5 .667 .286 .125 .056 .028 4 .429 .196 .095 .048 5 .571 .286 .143 .075 6 .393 .206 .111 7 .500 .278 .155 8 .607 .365 .210 9 .452 .274 10 .548 .545 11 .421 12 .500 15 .579 n = 6 m U 1 2 5 4 5 6 0 a45 .056 .012 .005 .002 .001 1 .286 .071 .024 .010 .004 .002 2 .428 .145 .048 .019 .009 .004 5 . .571 .214 .085 .055 .015 .008 4 .521 .151 .057 .026 .015 5 .429 .190 .086 .041 .021 6 .571 .274 .129 .065 .052 7 .557 .176 .089 .047 .8 .452 .258 .125 .066 9 .548 .505 .165 .090 10 .581 .214 .120 11 .457 .268 .155 12 .545 .551 .197 15 .596 .242 14 .465 .294 15 .555 .550 16 .409 17 .469 18 .551 Table IV. n = 7 \ 1 2 5 4 5 6. 7 u\ 0 .125 J028 .008 .005 .001 .001 .000 1 .250 .056 ..017 .006 .005 .001 .001 2 .575 .111 .055 .012 .005 .002 .001 5 .500 .167 .058 .021 .009 .004 .002 4 .625 .250 ..092 .056 .015 .007 .005 5 .555 .155 .055 .024 .011 .006 6 .444 .192 .082 .J057 .017 .009 7 .556 .258 .115 .055 .026 .015 8 .535 .158 .074 .057 .019 -S .417 •206 .101 .051 .027 10 .500 .264 .134 .069 .036 U .585 .524 .172 .090 .049 12 .594 .216 .117 .064 15 .464 .265 .147 .082 , 14 .558 .519 .183 .104 15 .578 .223 .130 16 .458 .267 .159 17 .500 .314 .191 18 .562 .365 .228 19 .418 .267 i 20 .475 .310 21 .527 .555 22 .402 25 .451 24 .500 25 .5^9 Table V 321 I M i m i l l TO STVl'lSTIt AI. WAI.VSIS T Mil 1 10. ( un n VI , \ VI.1 ('I'vvo-tail i H ' r i ' i ' M t i i i c r [nmils 1 — i l l r f o r tin' lull 1 III I l i l l l H I I U I Sn.s f o r /< TI.SI \ i I ' i *»*; 10 ' i 2.V , V 1', 5', 10', 25', . 1 Hi | 13 15 Hi is 17 1 1 Hi 17 Hi 3 (I •IS 1 1 Hi 17 10 1 II T.l 15 17 IS io . " i II (1 .•ill 15 17 is 20 Ii (1 II 1 ,->l 15 is 10 20 7 (I ( i 1 Hi IS III 21 s 0 0 1 1 ;>3 Hi IS 20 21 !l II 1 1 2 51 17 10 20 ''2 III (1 1 1 2 55 17 j HI 20 22 1 1 II 1 2 3 .Mi 1 7 ! 20 21 23 12 1 2 2 3 57 IS 1 20 21 23 i : ( 1 2 3 3 58 IS 21 22 2 I 11 1 2 3 1 5«J 1-1 21 22 2 i Iii 2 3 a till Hi 21 23 ''5 III 2 3 i 5 I'll 20 22 23 25 17 2 1 i 5 02 20 22 2 1 25 IS 3 I ,"i i i l»3 20 23 2 1 20 Hi 3 i , r i I i l i l 21 23 21 20 2(1 3 .') •"> I i 05 21 21 25 27 21 I i ;» i i 7 lili 22 21 25 ! 27 i i i 7 07 22 25 20 28 2^ i i; 4 H liS 22 25 20 28 21 7 s till 23 25 27 20 2."i .") 7 7 \\ 70 23 20 27 20 2ti i i 7 s '.1 71 21 2(1 28 30 '27 ti 7 S 10 72 21 27 28 30 2s 11 s 11 10 73 25 27 28 31 20 7 8 II 10 71 25 28 20 31 30 7 0 in 11 75 25 28 20 32 31 7 i 0 i II 11 7i i 20 2S 30 32 32 8 '.I 10 12 77 20 20 30 32 33 S Ml 11 12 7s 27 20 31 33 li l It 10 11 13 7'.i 27 30 31 33 »."• !l . II 12 13 so 2S 30 32 31 ;n, '.l 1 1 12 1 1 SI 2S 31 32 31 :t7 111 12 13 1 1 S2 28 31 3:< 35 38 Hi 12 13 1 1 S3 20 :i2 3;( .15 30 II" 12 13 I'i SI 2'.i 32 3;> 30 IH 11 13 1 1 15 S.'i 30 32 31 ,'it, II II 13 I I Iii SI. :«> 33 :u 3" | J 12 1 1 l.'i H. s7 31 33 35 37 i:< 12 II l.*i 17 S S 31 31 35 :<s II 13 I'i It, 17 V I 31 31 31. 3s 1.1 13 1". Ii. I Is 'Ml 32 t 35 30 30 r vnli|i"> i f V hirncr III.III '.HI. :i > | l l i i \ i l l i : i t r \ i l l « 1 « o t r limy In I..IIM.11 >\ Ink Kciiri'st i n l . p r l<ss than i.Y - I 2 - k \ .V -r I, WIHTI- k i> 1.2870, U.08INI. 0.8221. 0 .5752 for tin- I, 5. 10, 2 5 ' , v: t lm». rr>|M'rtivrly. A P P E N D I X B.  CORTICAL KAFS AKD ABLATIONS Plate I Map of Monkey Brain (After Brodmann, I9O9), Plate II Map of Cat Brain (After Garol, 1°42 c). Plate III Cortical Maps. Plate IV Cortical areas assumed for ablations. Plate V Maps of ablations a. Primary parietal lesions. b. Primary frontal lesions. c. Primary frontal lesions. d. Primary temporal lesions. e. Primary temporal lesions. f. Discards P l a t e I . Plate II Plate III FIG. I LABELS IDENTIFY SULCI FIG. 2 COMPOSITE MAP OF FUNCTIONAL CORTI CE S P l a t e IV . 3 B O U N D A R I E S O F C O R T I C A L A R E A S T E M P O R A L ( I N C L U D I N G A U D I T O R Y A N D I N S U L A R C O R T I C E S ) P A R I E T A L ( N O N S P E C I F I C ) F R O N T A L P l a t e V. LIT P l a t e V:b.~ XLVII P l a t e V 9. P l a t e V e. X X X I V I P l a t e V f . THE APPARATUS P l a t e I" P l a t e I I P l a t e I I I P l a t e IV Fl a n of the maze. Top View. Side View. Waiting Cages. Plate I . SCALE ALL DIMENSIONS ARE IN CENTIMETERS P L A N OF "Y" M A Z E 

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