Organization of a program of automatized food-procuring movement in rats

With the use of techniques of video- and photorecording, local destruction of the brain structures, and recording of the spike responses, roles of the suprasegmental brain structures in organization and control of automatized food-procuring movements were studied in rats under conditions of unrestrained behavior. It is shown that there are several neuronal populations in the prefrontal cortex related to programing, initiation, and current control of the efficacy of performance of food-procuring movements. The lateral hypothalamus is considered a motivational structure. It acts as the source providing an increase in the excitability and further support of the excitation level in the structures, which are links of the systems controlling food-procuring movements. The lateral hypothalamus is also involved in a system providing formation and final application of motor programs.     

The organization of instrumental food-procuring movements in rats]

The roles of the lateral hypothalamus, basolateral nucleus of the amygdalar complex, the second field of the frontal cortex, and ventromedial thalamic nucleus in organization of the fast ballistic food-procuring movements were studied in albino rats. Sequences of uni- and bilateral destruction of the brain structures were assessed by photorecording. Movement-related neuronal activity in these structures was recorded in freely moving animals. A specific involvement of each of the above listed structures in organization of food-procuring movements was shown. The lateral hypothalamus seems to participate in initiation of the motor program and its efficient realization, the basolateral amygdala appears to produce activatory, training, and stabilizing effects. The second field of the motor cortex leads in movement acquisition (i.e., in memorizing) and decision making about triggering the program. The idea about the relay role of the thalamic motor nucleus is supplemented by understanding of its more complex integrative function.     

Electrical activity of the lateral hypothalamus related to food-procuring movements of rats

On freely moving albino rats we demonstrated that, when fast food-procuring movements are performed, the mass electrical activity of the lateral hypothalamus (LH) is suppressed 1.6–2.0 sec before the movement beginning recorded with a photoelectrical device. Videorecording of the movements and recording of the spike activity of LH units showed that the latter are activated 1.0–0.1 sec before the movement initiation. The LH is considered a motivation-related structure, which serves as a source providing an increase in the excitability of the structures involved in the control of food-procuring movements and, further on, supporting this increased excitability. The LH is also a component of the mechanisms providing formation of the motor program. The role of the LH in the ensemble of motor centers, which organize and control voluntary movements, is discussed.     

Differences in hypothalamic monoaminergic mechanisms in heterogeneous conditioned reflexes in rats

Monoamines and aminoacids were microinjected in the middle medial hypothalamus of rats following the elaboration of conditioned food-procuring or avoidance reflexes. The glutamine acid had no effect on the avoidance reflex (AR), but stimulated the motor activity and the food-procuring reflex (FR), shortening its latencies, increasing the number of the conditioned food-procuring movements and the reflex size. Dopamine decreased the motor activity, but did not influence the AR latencies, whereas norepinephrine prolonged the AR latencies without changing the motor activity. Both catecholamines, while having no effect on the muscle tone, suppressed FR, prolonging the latencies and decreasing the reflex and the number of conditioned food-procuring movements. Local administration into the hypothalamus of GABA prolonged the latencies of heteromodal conditioned reflexes, whereas the serotonin had no effect on the recorded FR parameters, but facilitated AR significantly reducing its latencies.     

Disruption of food-getting movements after removal of the motor or premotor zone of the cerebral cortex in cats]

Instrumental food-procuring movements were studied in cats before and after unilateral or bilateral ablation of the motor or premotor cortical area. It is shown that unilateral impairment of the motor area affects the strength and accuracy of movements of the contralateral fore-leg, whereas the ablation of the premotor area leads to a slowing down of movements and breaking of a goal-directed movement into separate components. Bilataral ablation of the motor area irreversibly abolished the instrumental reflex. The ablation of the premotor cortex destroyed the animal's reaction to the sound signal, but food-procuring movements of the fore-legs were disturbed only temporarily. The obtained data are discussed on the basis of the concept that in cats the above cortical areas play different roles in the organization of goal directed behaviour.     

Roles of the Caudate Nuclei, Cerebellum, and Caudato-Cerebellar Interconnections in the Control and Coordination of Ballistic Food-Procuring Movements

We studied the roles of the cerebellum and caudate nuclei in the programming and control of ballistic movements. An experimental model of operant food-procuring movements of the rats was used; the activity of single neurons of the above structures was recorded in the course of these motor performances. To evaluate the impact of the cerebellar–caudate interaction on the process of control of the ballistic (centrally programmed) components of food-procuring motor performance, we also recorded modifications of the neuronal activity in one of the above-mentioned structures induced by electrical extrastimulation of another structure in the course of realization of the above components. It is demonstrated that the cerebellum and, in particular, its dentate nuclei are involved in the programming of ballistic food-procuring movements. Neurons of the caudate nuclei play a significant role in the immediate preparation for and subsequent current control of stereotyped ballistic movements. The high plastic properties of the cerebellar neurons manifested in the process of control of ballistic food-procuring movements are proved.     

A Functional Magnetic Resonance Imaging Study of Pathophysiological Changes Responsible for Mirror Movements in Parkinson’s Disease

Mirror movements correspond to involuntary movements observed in the limb contralateral to the one performing voluntary movement. They can be observed in Parkinson’s disease (PD) but their pathophysiology remains unclear. The present study aims at identifying their neural correlates in PD using functional magnetic resonance imaging. Ten control subjects and 14-off drug patients with asymmetrical right-sided PD were included (8 with left-sided mirror movements during right-hand movements, and 6 without mirror movements). Between-group comparisons of BOLD signal were performed during right-hand movements and at rest (p<0.005 uncorrected). The comparison between PD patients with and without mirror movements showed that mirror movements were associated with an overactivation of the insula, precuneus/posterior cingulate cortex bilaterally and of the left inferior frontal cortex and with a deactivation of the right dorsolateral prefrontal cortex, medial prefrontal cortex, and pre-supplementary motor area and occipital cortex. These data suggest that mirror movements in Parkinson’s disease are promoted by: 1- a deactivation of the non-mirroring inhibitory network (dorsolateral prefrontal cortex, pre-supplementary motor area); 2- an overactivation of prokinetic areas (notably the insula). The concomitant overactivation of a proactive inhibitory network (including the posterior cingulate cortex and precuneus) could reflect a compensatory inhibition of mirror movements.     

Automated Food-Procuring Movement-Related Neuronal Activity in the Basolateral Amygdala of the Rat

We studied the impulse activity of neurons of the basal and lateral amygdalar nuclei generated when experimental animals (rats) performed fast stereotyped food-procuring movements by the forelimb. Within the basolateral amygdala, there are neurons whose activity is related to different stages of getting off the food, and according to the characteristics of their spiking these neurons should be divided into a number of subpopulations. Activation forestalling the movement initiation by 0.5-1.0 sec was observed in most neurons of the basolateral amygdala; this is considered a manifestation of excitation related to a motivation component of the food-procuring behavior. Activation of amygdalar neurons following movement initiation can result from generation in this structure of additional excitation necessary for successful performance of a complete food-procuring motor cycle.     

The effect of ablation of the caudate nucleus on the alimentary zone of the lateral hypothalamus]

A conditioned food-procuring reaction previously elaborated to an acoustic stimulus was reproduced in chronic experiments on six cats by means of direct electrical stimulation of the posterior parts of the lateral hypothalamus. Folloiwng extensive bilateral electrolytic ablation of the caudate nucleus, conditioned food-procuring reaction to the stimulation of the hypothalamus could not be reproduced for 40 to 70 days. The conditioned foor-procuring reflex to the acoustic stimulus disappeared for 14 to 30 days to be subsequently spontaneously restored. After caudatotomy, a diminution of the average amplitude of background oscillations and of evoked potentials to acoustic stimuli was recorced in the examined zones of the lateral hypothalamus. The part played by the caudate nucleus in the processes of alimentary behaviour activation is discussed.     

Activity in the lateral prefrontal cortex reflects multiple steps of future events in action plans

To achieve a behavioral goal in a complex environment, we must plan multiple steps of motor behavior. On planning a series of actions, we anticipate future events that will occur as a result of each action and mentally organize the temporal sequence of events. To investigate the involvement of the lateral prefrontal cortex (PFC) in such multistep planning, we examined neuronal activity in the PFC of monkeys performing a maze task that required the planning of stepwise cursor movements to reach a goal. During the preparatory period, PFC neurons reflected each of all forthcoming cursor movements, rather than arm movements. In contrast, in the primary motor cortex, most neuronal activity reflected arm movements but little of cursor movements during the preparatory period, as well as during movement execution. Our data suggest that the PFC is involved primarily in planning multiple future events that occur as a consequence of behavioral actions.     

Lateral prefrontal cortex: architectonic and functional organization

A comparison of the architecture of the human prefrontal cortex with that of the macaque monkey showed a very similar architectonic organization in these two primate species. There is no doubt that the prefrontal cortical areas of the human brain have undergone considerable development, but it is equally clear that the basic architectonic organization is the same in the two species. Thus, a comparative approach to the study of the functional organization of the primate prefrontal cortex is more likely to reveal the essential aspects of the various complex control processes that are the domain of frontal function. The lateral frontal cortex appears to be functionally organized along both a rostral-caudal axis and a dorsal-ventral axis. The most caudal frontal region, the motor region on the precentral gyrus, is involved in fine motor control and direct sensorimotor mappings, whereas the caudal lateral prefrontal region is involved in higher order control processes that regulate the selection among multiple competing responses and stimuli based on conditional operations. Further rostrally, the mid-lateral prefrontal region plays an even more abstract role in cognitive control. The mid-lateral prefrontal region is itself organized along a dorsal-ventral axis of organization, with the mid-dorsolateral prefrontal cortex being involved in the monitoring of information in working memory and the mid-ventrolateral prefrontal region being involved in active judgments on information held in posterior cortical association regions that are necessary for active retrieval and encoding of information.     

Neuronal reactions in field 2 of the frontal cortex, related to organization and performance of food-procuring movements in rats

Movement-related electrical reactions of neuronal units localized in field 2 of the frontal cortex were studied in albino rats performing fast food-procuring movements under conditions of unrestrained behavior. According to the temporal characteristics of the changes in the neuronal spike activity, three types of reactions were classified: (i) activation that forestalled the movement initiation for 1.0–1.5 sec; (ii) activation or inhibition forestalling this beginning for 0.20–0.26 sec; and (iii) activation in the course of a performed movement. Considerations about the involvement of the neurons of various cortical layers in the mechanisms of programing, switching on, and current control of the efficiency of performance of food-procuring movements are proposed, and the role of the frontal cortex in these processes is discussed.     

Role of frontal cortex in the organization of rapid ballistic movements of rats

We have studied the characteristics of rapid ballistic food-procuring movements in nonpedigree albino rats and have established that after ablation of the second area of the frontal cortex contralaterally to the preferred extremity the number of attempts increased, the duration of the movements decreased, and the phase structure of the movements was reorganized. After bilateral ablation of the cortex the animals completely lost their skill at procuring food. Our results indicate involvement of the frontal cortex in the development and achievement of the motor programs produced.N. I. Pirogov Medical Institute. Ukrainian Ministry of Public Health. Translated from Neirofiziologiya, Vol. 24, No. 2, pp. 186–192, February, 1992.     

Top-down control of motor cortex ensembles by dorsomedial prefrontal cortex

Dorsomedial prefrontal cortex is critical for the temporal control of behavior. Dorsomedial prefrontal cortex might alter neuronal activity in areas such as motor cortex to inhibit temporally inappropriate responses. We tested this hypothesis by recording from neuronal ensembles in rodent dorsomedial prefrontal cortex during a delayed-response task. One-third of dorsomedial prefrontal neurons were significantly modulated during the delay period. The activity of many of these neurons was predictive of premature responding. We then reversibly inactivated dorsomedial prefrontal cortex while recording ensemble activity in motor cortex. Inactivation of dorsomedial prefrontal cortex reduced delay-related firing, but not response-related firing, in motor cortex. Finally, we made simultaneous recordings in dorsomedial prefrontal cortex and motor cortex and found strong delay-related temporal correlations between neurons in the two cortical areas. These data suggest that functional interactions between dorsomedial prefrontal cortex and motor cortex might serve as a top-down control signal that inhibits inappropriate responding.     

Characteristics of the systemic organization of the cortical neurons in various forms of the manifestation of the conditioned reflex to time

Conditioned food-procuring reflex to time (2 minutes interval) was elaborated in cats. By the method of cross-correlative analysis the combined neurones activity in microareas and between microareas of the motor cortex was compared at various forms of conditioned reflex manifestation. Three types of reactions were considered: A--decrease of respiratory movements amplitude towards the end of the studied interval between reinforcement; B--increase of the amplitude; C--food-procuring paw movement a few seconds before the reinforcement. All three forms of the conditioned reflex to time differed from each other by the level of increase of functional connections number by the moment of the reinforcing stimulus action, and also to a greater extent by the frequency of occurrence of intervals in which the sum of neuronal connections in all simultaneously recorded microareas was greater in the "active" phase than in the "inactive" one. These parameters did not always correlate with the change of impulses frequency of separate neurones which occurred considerably more seldom. All the observed changes were manifest significantly more often when the animals performed the instrumental food-procuring movement than during changes of only the respiratory movements amplitude.     

Dynamics of caudate nucleus involvement in the process of internal inhibition of the food-getting reflex]

Changes of slow spindle electrical activity (12 to 14 cps) and evoked potentials in the cat caudate nuclei and the motor cortex were studied at different stages of elaboration of differentiation inhibition of a conditioned food-procuring reflex to acoustic stimuli. It has been shown that participation of the caudate nuclei in the inhibitory process begins at the stage of enhancement of food-procuring activity (second stage) and is apparently due to the animal's strong alimentary motivational excitation. At the third stage the inhibitory influence of caudate nuclei on the animal's motor activity in general, and the activity of the motor cortex in particular, develops to the full.     

Neurochemical mechanisms of the septum in food getting conditioned responses in rats]

The influence of microinjections of monoamines and of glutamic acid into the lateral zone of the septum on a food-procuring conditioned reaction was studied in experiments on rats. Dopamine does not change, while noradrenaline, serotonin and glutamic acid increase the magnitude and number of conditioned food-procuring movements and substantially lengthen the reflex latency. The effects of serotonin are achieved through D-serotonin-, and of noradrenaline, through L-adrenoreactive structures of the neuronal systems of the septum. The influence of glutamic acid on the rat food-procuring conditioned reactions is not linked with the action on the serotonin- or adrenoreactive structures of the septum neuronal systems.     

The role of the lateral and medial hypothalamus in reproducing a motor reaction as the signal during the acquisition of classical conditioned reflexes]

In experiments on three dogs there was shown that testing electrostimulation of the lateral hypothalamus reproduced the motor reaction which is a signal stimulus at elaboration of classic alimentary conditioned reflexes (CRs) and did not reproduce it at elaboration of classic defensive CR. Testing electrostimulation of medial parts of the hypothalamus reproduced, as LH electrostimulation the "signal" motor reaction, but in less percentage of cases, during elaboration of classic alimentary CRs and did not reproduce it at elaboration of classic defensive CRs. The reproduction of the signal motor reaction at LH electrostimulation is connected with activation of backward conditioned connection from motivation structures of the hypothalamus to representation of the signal stimulus in the motor cortex.     

Competing neural responses for auditory and visual decisions

Why is it hard to divide attention between dissimilar activities, such as reading and listening to a conversation? We used functional magnetic resonance imaging (fMRI) to study interference between simple auditory and visual decisions, independently of motor competition. Overlapping activity for auditory and visual tasks performed in isolation was found in lateral prefrontal regions, middle temporal cortex and parietal cortex. When the visual stimulus occurred during the processing of the tone, its activation in prefrontal and middle temporal cortex was suppressed. Additionally, reduced activity was seen in modality-specific visual cortex. These results paralleled impaired awareness of the visual event. Even without competing motor responses, a simple auditory decision interferes with visual processing on different neural levels, including prefrontal cortex, middle temporal cortex and visual regions.     

Frontal lobe and cognitive development

In phylogeny as in ontogeny, the association cortex of the frontal lobe, also known as the prefrontal cortex, is a late-developing region of the neocortex. It is also one of the cortical regions to undergo the greatest expansion in the course of both evolution and individual maturation. In the human adult, the prefrontal cortex constitutes as much as nearly one-third of the totality of the neocortex. The protracted, relatively large, development of the prefrontal cortex is manifest in gross morphology as well as fine structure. In the developing individual, its late maturation is made most apparent by the late myelination of its axonal connections. This and other indices of morphological development of the prefrontal cortex correlate with the development of cognitive functions that neuropsychological studies in animals and humans have ascribed to this cortex. In broad outline, the ventromedial areas of the prefrontal cortex, which with respect to otherprefrontal areas develop relatively early, are involved in the expression and control of emotional and instinctual behaviors. On the other hand, the late maturing areas of the lateral prefrontal convexity are principally involved in higher executive functions. The most general executive function of the lateral prefrontal cortex is the temporal organization of goal-directed actions in the domains of behavior, cognition, and language. In all three domains, that global function is supported by a fundamental role of the lateral prefrontal cortex in temporal integration, that is, the integration of temporally discontinuous percepts and neural inputs into coherent structures of action. Temporal integration is in turn served by at least three cognitive functions of somewhat different prefrontal topography: working memory, preparatory set, and inhibitory control. These functions engage the prefrontal cortex in interactive cooperation with other neocortical regions. The development of language epitomizes the development of temporal integrative cognitive functions and their underlying neural substrate, notably the lateral prefrontal cortex and other late-developing cortical regions.