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THE TWO COMPONENTS of the GRASP REFLEX AFTER ABLATION of FRONTAL CORTEX IN,MONKEYS* by GEOFFREY Rushwortht and D

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THE TWO COMPONENTS of the GRASP REFLEX AFTER ABLATION of FRONTAL CORTEX IN,MONKEYS* by GEOFFREY Rushwortht and D J Neurol Neurosurg Psychiatry: first published as 10.1136/jnnp.22.2.91 on 1 May 1959. Downloaded from J. Neurol. Neurosurg. Psychiat., 1959, 22, 91. THE TWO COMPONENTS OF THE GRASP REFLEX AFTER ABLATION OF FRONTAL CORTEX IN,MONKEYS* BY GEOFFREY RUSHWORTHt and D. DENNY-BROWN From the Neurological Unit, Boston City Hospital, Boston, Mass., U.S.A., and the Department of Neurology, Harvard Medical School A grasp reflex was shown by Richter and Hines of the upper limb during application of the stimulus (1932) to result from experimental lesions in the to the palm leads one to suppose that shoulder pre-motor region (which they identified with area traction had not been recognized as a potent stimulus, six of Brodmann) of the frontal lobes in adult of itself, of reflex closure of the fingers (the traction monkeys. Reflex grasping in the monkey was later response). studied in more detail by Fulton and his associates In this paper we shall refer to the grasp reflex as in an effort to relate its occurrence to ablation of the finger flexion, with thumb adduction that occurs specific frontal areas and to analyse its nature at the in response to a distally moving pressure contact periphery (Bieber and Fulton, 1933, 1938; Kennard applied to the palm preceding traction on the finger and Fulton, 1933; Fulton, 1934). Since Janischew- flexors, with the shoulder, arm, and forearm fixed. Protected by copyright. sky (1909) first described reflex hand grasping in a It is regularly present after frontal lobe lesions in the patient with organic brain disease, it has been the monkey. It is to be compared with the "instinctive subject of many reports and several studies. Walshe grasp reaction" which is the response to a light and Robertson (1933) and Walshe and Hunt (1936) stationary or moving tactile stimulus on any part of regarded the human grasp reflex as a purely pro- the hand and it consists of orientation of the hand prioceptive reflex being devoid of visual or tactile into a position necessary for finger-thumb palpation elements, and a similar view of the simian grasp and final grasping of the object. This reaction, reflex was reached by Bieber and Fulton (1938). which is independent of vision, is occasionally found Seyffarth and Denny-Brown (1948) came to different in the normal monkey, though the more usual conclusions for the human grasp reflex when they response to a light tactile stimulus to the hand is one found that the adequate stimulus for it was a of slight withdrawal (avoiding). The instinctive distally moving stimulus to certain areas of the palm. grasp reaction is released by frontal lobe lesions in They were able to dissociate this triggering effect the monkey. Finally, the traction response has to be from the proprioceptive element. An essential clearly differentiated from the other two grasping sensory input for the reflex was derived from the automatisms. Traction response is reflex finger skin and subcutaneous tissue of the palm, and this flexion and thumb adduction resulting from abduc- http://jnnp.bmj.com/ sensitized a secondary proprioceptive component tion and flexion or traction on the shoulder, and it when the stimulus stretched the long finger flexors. can also reinforce the grasping automatisms outlined This secondary component reinforced and main- above. It is a synergistic effect of stretching the tained the closing phase of the grasp initiated by spastic muscles at the shoulder and it is present palmar stimulation. therefore in the thalamic monkey as Bieber and The large literature on reflex grasping shows many Fulton (1938) described. It is unaffected by deaffer- apparent discrepancies of observation and inter- entation of the hand and arm, but not the shoulder pretation which are doubtless due to the confusion (Twitchell, 1954) and is modified, as are stretch on October 1, 2021 by guest. of terminology, lack of clear definition, and the reflexes, by head and neck turning. Though it is not failure to differentiate three distinct grasping auto- a true grasp reflex (as defined above), an animal matisms (Seyffarth and Denny-Brown, 1948). In rendered spastic experimentally and showing the particular the lack of information regarding fixation traction response can learn to use it for a hooking type of grasping such as hanging from a bar. Later *Assisted by a grant from the Harrington Fund. it may learn to project the arm to initiate the traction tOn leave ofabsence from the University Laboratory of Physiology, response of flexion of the fingers which can then be Oxford. Present address: Neurological Research Unit, The Churchill Hospital, Oxford. hooked around an object. The subsequent slight 91 J Neurol Neurosurg Psychiatry: first published as 10.1136/jnnp.22.2.91 on 1 May 1959. Downloaded from 92 GEOFFREY RUSHWORTH AND D. DENNY-BROWN stretching of the spastic finger flexors by the object may then result in closing the fingers through a facilitated stretch reflex and so lead to a fixed resistance to opening of the hand. It is therefore necessary to distinguish the purely passive traction response from grasping as a positive act. The present experiments were designed to clarify the grasp reflex of monkeys after frontal lobe ablation and to demonstrate the part played by the efferent muscle spindle innervation (gamma motor fibres) in this reflex. Methods Monkeys (M. cyamologous) weighing between 2,200 and 3,000 g. were employed in this study. Their behaviour was watched for several days before operation and they COR 34 were examined neurologically while strapped in a small COR 35 chair designed to leave the limbs free while restraining ---- COR 39 the body. Under sterile conditions and nembutal oo** COR40 anaesthesia (administered by the intraperitoneal route), the skull was opened by turning a left-sided bone flap. FIG. 1.-The extent of ablation of the left frontal lobe in the four monkeys described in the text, charted from the necropsy The central sulcus and the anatomical features of areas specimens. Where the ablation was bilateral it was approximately 4, 6, and 8 of Brodmann were identified. The left frontal symmetrical. In animals COR 34 and COR 35 the entire cortex pole was amputated, and areas 8 and 6 removed by in front of the precentral sulcus was removed, whereas in animals COR 39 and COR 40 the precentral gyrus remained. C.S. is the subpial suction through small holes in the pia-arachnoid. Protected by copyright. The pia collapsed over the ablated areas. Much attention central sulcus. was given to the orbital surface and the cingulate (area 24) gyrus. The former was removed as far back as selves. Finger jerks, response to pin prick in the palm, possible and the cingulate to the level of the medial limit effects of turning the head and neck, and spontaneous of the central sulcus. After haemostasis had been secured, struggling movements could be recorded. the dura was carefully repaired, the bone flap replaced About six weeks after the first operation, the procedure and fixed by sutures in the galea. The muscles were was repeated in the right side of the head and the grasp closed in layers and the skin re-approximated by con- reflex studied in both hands. The behaviour of the tinuous subcutaneous silk suture. animals after frontal lobe lesions is to be reported at a By the next day the animals were usually quite active later date. The extent of the ablations in the four and at the end of a week the wound had healed sufficiently animals discussed in this paper is shown in Fig. 1, and is to allow of catching the animal in a net and strapping it drawn from the necropsy specimens which had been into a special chair for neurological examination and fixed in situ. experiment. The chair was provided with a wooden arm projecting at right angles to the chair-back and the Results animal's right forearm was fixed to this by strips of Within one week of operation a grasp reflex was "elastoplast" around the elbow and wrist. A light regularly elicitable from the palm contralateral to the rubber band was fixed to the finger tips and attached to a cortical lesion and more facile avoiding responseshttp://jnnp.bmj.com/ Grass strain-gauge myograph with sufficient tension to from the homolateral limbs ("transcortical release"). keep the fingers just extended. A small concentric needle When the animal was strapped into the experimental electrode was placed within the flexor muscles of the a fingers and lightly fixed there. The potentials picked up chair and its forearm fixed to the chair arm, grasp by this and the output of the strain-gauge were suitably reflex was still elicitable, though greater pressure of amplified and displayed on a cathode ray tube which was the moving object was now necessary to elicit the photographed on moving paper by means of a Grass reflex every time. It was also necessary to see that camera. The stimulus for the grasp reflex was provided the dorsum of the fingers and hand did not make by a small insulated metal strip fitted with electrical contact with any part of the experimental chair, on October 1, 2021 by guest. contacts. The strip was moved with moderate pressure because tactile stimuli to the dorsum of the hand are across the palm beginning proximally and the initial powerful inhibitors of the grasp reflex. Neither the contact with the palm was signalled automatically as of the to a a was the increased resistance to movement as the stimu- attaching fingers myograph through lator encountered the flexing fingers. The grasp reflex so light rubber band nor the insertion of a small co- elicited was studied before and after injection of 1 % axial electrode into the finger flexors had any signi- procaine around the ulnar and median nerves at the wrist, ficant effect on the grasp reflex.
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