**5. Anterior cingulate circuit**

The anterior cingulate circuit (limbic loop) is important in behavior control and adaptation of behaviours after making a mistake (ibid.). The damage of this circuit results in emotional disorders especially deep apathy and lack of spontaneity. Lowered mood is accompanied by weakening of affect and motor adynamy (ibid.).

On the basis of a pattern of basal ganglia connections, being a part of particular loops with cerebral cortex, similarity of motor and emotional functions can be deducted (Alexander et all., 1990). Information processed by various loops partly overlap in the striatum (ibid.). In the globus pallidus pars internalis and the substantia nigra pars reticularis and subsequently in the thalamus pieces of information from various circuits converge. Pieces of information from the thalamus reach limited areas of the cerebral cortex, which control motor, emotional and cognitive functions (ibid). Such functional organization makes it possible to select motor and mental actions depending on information incoming from external and internal environment (Mink, 1999; Morgane et all., 2005; Groenewegen & Dongen, 2007).

The circuits, described above, functionally connect the basal ganglia with the cerebral cortex (Alexander et all., 1986; DeLong et all., 1998; Elliot, 2003; Haber, 2003; Saint-Cyr et all., 2003; Morgane et all., 2005; Olzak & Gorzelańczyk, 2005; Groenewegen & van Dongen, 2007; Laskowska et all., 2008; Haber et all., 2009). The assumption that the circuits connecting the cerebral cortex with the basal ganglia work independently and in a parallel way (Alexander et all., 1986, 1990; DeLong et all., 1998) has expired. Various consequences of the damages of particular loops activity depend on the cerebral cortex areas with which they connect and on the circumstances in which they are activated. The degree of co-operation of particular basalo-thalamo-cortical loops is not known (Longstaff, 2006; Laskowska et all., 2008; Haber et all., 2009; Sadikot et all., 2009), however more and more data indicate that the exchange of information between particular circuits takes place (McFarland et all., 2002; Groenewegen & van Dongen, 2007; Laskowska et all., 2008; Haber & et all., 2009; Sadikot et all., 2009).

Subcortical nuclei relate not only to motor control, but also to the processes of reminding as well as executive functions processes, short-term memory, the analysis of mutual setting of objects and to undertaking actions (Sławek et all., 2001; Frank et all., 2001; Royall et all., 2002; Elliot, 2003; Haber, 2003; Laskowska et all., 2008; McNab et all., 2008; Haber et all., 2009).

The basal ganglia take part in the control of motor, emotional and cognitive behaviours by two pathways (indirect and direct) exerting contradictory effect on the stimulation of the thalamus and the cerebral cortex. (Albin et all., 1989; DeLong, 1990; Obeso et all, 1997, 2000b; Sławek, 2003; Sobstyl et all., 2003; Groenewegen, 2003; Longstaff, 2006; DeLong & Wichmann, 2007; Groenewegen & van Dongen, 2007; Szołna, 2007). In the nigrostriatal pathway there are two kinds of dopaminergic receptors relating to striatal medium spiny neurons: D1, which activate GABA-ergic neurons of the putamen in the direct pathway; and D2, whose stimulation inhibits GABA-ergic neurons of the putamen in the indirect pathway (ibid.).

The direct pathway relates to striatal medium spiny neurons (SP/DYN) releasing GABA and inhibiting GABA-ergic neurons running from GPi and SNpr to the thalamus. Cortical stimulation of this pathway causes activation of thalamus neurons (Groenewegen, 2003; Sobstyl et all., 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007; Szołna, 2007). The direct pathway connects the putamen, where the connections from the cerebral cortex and the substantia nigra (SNpc) meet, with the exit structures - the internal part of the globus pallidus and reticular part of substantia nigra pars reticulata – (SNpr) (ibid.).

The anterior cingulate circuit (limbic loop) is important in behavior control and adaptation of behaviours after making a mistake (ibid.). The damage of this circuit results in emotional disorders especially deep apathy and lack of spontaneity. Lowered mood is accompanied by

On the basis of a pattern of basal ganglia connections, being a part of particular loops with cerebral cortex, similarity of motor and emotional functions can be deducted (Alexander et all., 1990). Information processed by various loops partly overlap in the striatum (ibid.). In the globus pallidus pars internalis and the substantia nigra pars reticularis and subsequently in the thalamus pieces of information from various circuits converge. Pieces of information from the thalamus reach limited areas of the cerebral cortex, which control motor, emotional and cognitive functions (ibid). Such functional organization makes it possible to select motor and mental actions depending on information incoming from external and internal

The circuits, described above, functionally connect the basal ganglia with the cerebral cortex (Alexander et all., 1986; DeLong et all., 1998; Elliot, 2003; Haber, 2003; Saint-Cyr et all., 2003; Morgane et all., 2005; Olzak & Gorzelańczyk, 2005; Groenewegen & van Dongen, 2007; Laskowska et all., 2008; Haber et all., 2009). The assumption that the circuits connecting the cerebral cortex with the basal ganglia work independently and in a parallel way (Alexander et all., 1986, 1990; DeLong et all., 1998) has expired. Various consequences of the damages of particular loops activity depend on the cerebral cortex areas with which they connect and on the circumstances in which they are activated. The degree of co-operation of particular basalo-thalamo-cortical loops is not known (Longstaff, 2006; Laskowska et all., 2008; Haber et all., 2009; Sadikot et all., 2009), however more and more data indicate that the exchange of information between particular circuits takes place (McFarland et all., 2002; Groenewegen &

environment (Mink, 1999; Morgane et all., 2005; Groenewegen & Dongen, 2007).

van Dongen, 2007; Laskowska et all., 2008; Haber & et all., 2009; Sadikot et all., 2009).

Subcortical nuclei relate not only to motor control, but also to the processes of reminding as well as executive functions processes, short-term memory, the analysis of mutual setting of objects and to undertaking actions (Sławek et all., 2001; Frank et all., 2001; Royall et all., 2002; Elliot, 2003; Haber, 2003; Laskowska et all., 2008; McNab et all., 2008; Haber et all., 2009). The basal ganglia take part in the control of motor, emotional and cognitive behaviours by two pathways (indirect and direct) exerting contradictory effect on the stimulation of the thalamus and the cerebral cortex. (Albin et all., 1989; DeLong, 1990; Obeso et all, 1997, 2000b; Sławek, 2003; Sobstyl et all., 2003; Groenewegen, 2003; Longstaff, 2006; DeLong & Wichmann, 2007; Groenewegen & van Dongen, 2007; Szołna, 2007). In the nigrostriatal pathway there are two kinds of dopaminergic receptors relating to striatal medium spiny neurons: D1, which activate GABA-ergic neurons of the putamen in the direct pathway; and D2, whose stimulation inhibits GABA-ergic neurons of the putamen in the indirect pathway

The direct pathway relates to striatal medium spiny neurons (SP/DYN) releasing GABA and inhibiting GABA-ergic neurons running from GPi and SNpr to the thalamus. Cortical stimulation of this pathway causes activation of thalamus neurons (Groenewegen, 2003; Sobstyl et all., 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007; Szołna, 2007). The direct pathway connects the putamen, where the connections from the cerebral cortex and the substantia nigra (SNpc) meet, with the exit structures - the internal part of the globus

pallidus and reticular part of substantia nigra pars reticulata – (SNpr) (ibid.).

**5. Anterior cingulate circuit** 

(ibid.).

weakening of affect and motor adynamy (ibid.).

The direct pathway is connected with striatal medium spiny neurons (ENK) secreting GABA and it leads through the external part of the globus pallidus (GPe) to STN, which stimulates inhibitory neurons in GPi and SNr structures. Cortico-striatal stimulation of the indirect pathway decreases thalamus stimulations and inhibits motor cerebral cortex (Albin et all., 1989; DeLong, 1990; Obeso et all., 1997, 2000b; Herrero et all., 2002; Sobstyl et all., 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007).

Dopamine is a neurotransmitter in SNc neurons (Groenewegen, 2003; Sobstyl et all., 2003; Groenewegen & van Dongen, 2007). During repose occasional discharges of small frequency, which do not affect movement, appear in these neurons (Longstaff, 2006). The frequency of neurons firing changes when a stimulus rewarding a movement is present (ibid.). They influence the reply of striatal medium spiny neurons to the stimulation from the cerebral cortex. GABA/SP/DYN neurons increase responsiveness in the direct pathway, however GABA/ENK neurons reduce responsiveness in the indirect pathway due to the SNc influence. As a result, the nigro-striatal connection increases the activity of the direct pathway, inhibiting the indirect pathway (ibid.).

The basal ganglia are responsible, inter alia, for executing motor sequences (Alexander et all., 1990; Mink, 1999; Sadowski, 2001; Herrero et all., 2002; McFarland et all., 2002; Longstaff, 2006; Haber et all., 2009). Each sequence is represented by striatal medium spiny neurons creating motor or oculomotor microloops in the cortico-subcortical circuit, which can be activated or inhibited (Alexander et all., 1986; Longstaff, 2006). A considerable part of striatal medium spiny neurons has low frequency of electric discharge at rest (0,1-1Hz). In other subcortical nuclei - GPi and SNr, the frequency of electric discharge at rest is considerably higher and it amounts to about 100*Hz* (ibid.). According to a contemporary model of motor control carried out by the basal ganglia, movements are initiated by activation of the motor cerebral cortex, which consequently activates the striatum (Groenewegen, 2003; Sobstyl et all., 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007; Sadikot et all., 2009). The activity of striatal medium spiny neurons increases during the execution of movements being a result of functioning of cortico-striatal neurons (Longstaff, 2006). When a discharge frequency of GPi and SNr decreases, a reduction in the thalamus inhibition follows making it possible to execute a movement or other mental action (Albin et all., 1989; DeLong et all., 1990; Obeso et all., 1997, 2000; Sobstyl et all., 2003; Longstaff, 2006).

Motor, emotional and cognitive functions are controlled by the system of structures and connections of the central nervous system, among which three levels of co-operation were distinguished: lower, middle and upper (Schotland & Rymer, 1993; Longstaff, 2006). Experimentally induced damages of the nervous system centers in animals result in the lack of the function subjected to the place of the damage and the appearance of new or enhancement of until now executed actions. A damage of the motor center of the cerebral cortex causes limbs paresis, an increase in muscles tension, intensification of tendinous reflexes in these limbs as well as appearance of Babiński's reflex (Babinski, 1986). The occurrence of such symptoms indicates that lower in hierarchy structures are inhibited by higher ones and that the lack of this inhibition causes unblocking of the physiologically inhibited centre. Clinical observations of persons with the basal ganglia damages indicate that these structures do not initiate movements, but they control the course of skilled sequence of movements (ibid.). An increase in the activity of the direct pathway increases, and of the indirect pathway decreases, stimulation of particular areas of the cerebral cortex (Groenewegen, 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007; Szołna, 2007).

Functional Anatomy, Physiology and Clinical Aspects of Basal Ganglia 99

such as for example: making decisions, the choice of motor action, the change of behavior, working memory can be disturbed in a similar degree, independently from which structure of a particular loop is damaged (ibid.). Because of a specific structure of striatal medium spiny neurons (having a very large quantity of synaptic connections) and synaptic processes like long-term synaptic potentiation (LTP) and long-lasting synaptic depression (LTD), similar to those carried out in the cerebral cortex, the striatum (including the ventral striatum) is functionally the main structure of cortico-subcortical loops (Picconi et all., 2005) that controls motor, emotional and cognitive functions. Due to numerous inter-striatal and cortico-striatal connections and described above plastic mechanisms enabling enhancement or reduction of stimulations, the striatum together with the cerebral cortex controls the functions of particular cortico-subcortical loops, substantially influencing mental processes

The results obtained from the experiments on a model of shoulder movement simulation indicate that the higher dopamine concentration is in the striatum the shorter is the time necessary to initiate a movement (ibid.). This means that the time necessary to initiate a movement becomes shorter simultaneously with an increase of the striatum stimulation (ibid.). Moreover, the simulation of a decrease in dopamine concentration in the striatum in the same model causes bradykinesia and akinesia, similar to that observed in the Parkinson's disease (Prescott et all., 2002). The key function of the striatum is to produce signals which reach, through the thalamus, the cerebral cortex (ibid.). It was suggested that the choice of a particular pattern of action takes place in the striatum due to so called gating mechanism. Activation of the neurons responsible for processing of a particular pattern leads to inhibition of other striatal neurons. The mechanism makes it possible e.g. to process efficiently information introduced to the working memory with simultaneous inhibition of the inflow of new information before completing a presently executed task (Frank et all., 2001). If the gate is closed, new information does not influence the memory essentially and that is why it allows a stable maintenance of information being processed (ibid.). Opening of the gate enables data updating (ibid.). The inhibition of new information inflow prevents their interference with previously gathered data (ibid.). This mechanism enables continuous selection of the processed information optimizing in this way execution of mental actions (ibid.). The disturbance of the mechanism of selective actualization of data in proper time (both too frequent and too rare data updating) increases the number of errors made in various tasks. This disorder relates to initiation of both movement and thought processes. A confirmation of the concept equity is psychic akinesia observed in persons with a damage of the striatum (Brown and Marsden, 1990). However, inhibition of the internal part of the globus pallidus leads to disinhibition of frontal loops activity, which results in continuous updating of data. Continuous updating of data is caused by the lack of blocking in access to the working memory (Frank et all., 2001). Results of these disorders can be Absentmindedness, impulsivity and hyperactivity (occurings in the disorders such as: Huntington's disease, Tourette syndrome, attention deficit hyperactivity disorder) can be the results of the above mentioned disorders (ibid.). Diminishing of dopamine concentration in the striatum causes that the gate regulating the access to the working memory is opened and closed in the wrong time, and in permanent inhibition of the internal part of the globus

The gating model does not fully explain the selection of data, which arises from the fact that dopamine is secreted in large areas of the prefrontal cortex. In this model, different areas of the prefrontal cortex are activated simultaneously, making it impossible to select one

(Brown et all., 1997; Prescott et all., 2002).

pallidus it is opened all the time (ibid.).

Cooperation between the direct and indirect pathways changes the activity of the cerebral cortex in such a way that the activity increases in the areas which control execution of a particular motor action and inhibits the areas which are not involved in the executed activity (ibid.). Planning of motor activity relates to the functioning of the cerebral prefrontal cortex controlling the process of scheduling and execution of a particular sequence of movements (LeDoux, 1996; Groenewegen, 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007). The control of implementation of the actions scheduled is possible when the the centers of the cerebral cortex and basal ganglia (especially the caudate) cooperate properly (Alexander et all., 1986, LeDoux, 1996; Herrero et all., 2002; Groenewegen & van Dongen, 2007). The nucleus caudatus – a part of the ventral striatum is connected with a substantial part of the frontal, mainly with the associative cerebral cortex (ibid.). Pieces of information incoming from the cerebral cortex to the caudate make it possible to control and implement a particular sequence of movements (ibid.). A program relating to the implementation of a specific sequence of movements is sent from the caudate through the thalamus to the premotor prefrontal cerebral cortex, which controls voluntary movements (ibid.). The basal ganglia in connection with ventral nuclei of the thalamus and frontal areas of the cerebral cortex play an essential role in instrumental conditioning (Petri et all., 1994; Smith's Petri et all., 2006; Rueda-Orozco et all., 2008). The results obtained in investigations on animals confirm the meaning of the basal ganglia in the procedural memory (Petri et all., 1994; Akhmetelashvili et all., 2007; Hartman et all., 2008; Rueda-Orozco et all., 2008).

The pattern of basal-thalamic-cortical connections carrying out motor, emotional and cognitive functions is mutual for these functions (Alexander et all., 1996; DeLong et all., 1998; Herrero et all., 2002; Groenewegen, 2003; Saint-Cyr, 2003; Groenewegen & van Dongen, 2007; Laskowska et all., 2008), and the control of these functions consists in the choice of the sequence of behaviours adapted to a particular situational context (Heilman et all., 1993; Royall et all., 2002; Longstaff, 2006).

The control of motor, emotional and cognitive functions is hierarchical and the processing of information connected with these functions can be executed sequentially or simultaneously in various structures of the brain. Clinical observations and the results of the investigations carried out in persons with a damage of the basal ganglia confirm the participation of subcortical structures not only in motor but also cognitive functions (Oberg and Divac, 1979; Berns and Sejnowski, 1995; Frank et all., 2001). Also the results of computer simulations of cognitive actions such as: maintenance of information in operating memory, remembering, planning of sequences of behaviors and making decisions indicate a participation of the basal ganglia in the control of cognitive functions (Prescott et all., 2002; Jessup et all., 2011). The function of the basal ganglia in the control of motor, emotional and cognitive functions relates to cooperation with frontal cerebral cortex areas (Brown et all., 1997). The basal ganglia cooperating with the prefrontal cerebral cortex make it possible to initiate motor, emotional and cognitive actions. A complex system of stimulations (inhibition and disinhibition) limits the flow of excessive information which disables the initiation of an activity. A complex choice mechanism makes it possible to initiate a particular kind of movement without detailed determination of the whole sequence of the movements executed (Bullock & Grosberg, 1988; Hikosaka, 1989; Chevalier & Deniau, 1990; Passingham, 1993).

According to the concept of mental (motor, emotional, cognitive) functions control carried out by cortico-subcortical loops a proper action of all the structures of these loops is a condition necessary for their efficient functioning (Brown et all., 1997). Mental processes,

Cooperation between the direct and indirect pathways changes the activity of the cerebral cortex in such a way that the activity increases in the areas which control execution of a particular motor action and inhibits the areas which are not involved in the executed activity (ibid.). Planning of motor activity relates to the functioning of the cerebral prefrontal cortex controlling the process of scheduling and execution of a particular sequence of movements (LeDoux, 1996; Groenewegen, 2003; Longstaff, 2006; Groenewegen & van Dongen, 2007). The control of implementation of the actions scheduled is possible when the the centers of the cerebral cortex and basal ganglia (especially the caudate) cooperate properly (Alexander et all., 1986, LeDoux, 1996; Herrero et all., 2002; Groenewegen & van Dongen, 2007). The nucleus caudatus – a part of the ventral striatum is connected with a substantial part of the frontal, mainly with the associative cerebral cortex (ibid.). Pieces of information incoming from the cerebral cortex to the caudate make it possible to control and implement a particular sequence of movements (ibid.). A program relating to the implementation of a specific sequence of movements is sent from the caudate through the thalamus to the premotor prefrontal cerebral cortex, which controls voluntary movements (ibid.). The basal ganglia in connection with ventral nuclei of the thalamus and frontal areas of the cerebral cortex play an essential role in instrumental conditioning (Petri et all., 1994; Smith's Petri et all., 2006; Rueda-Orozco et all., 2008). The results obtained in investigations on animals confirm the meaning of the basal ganglia in the procedural memory (Petri et all., 1994;

Akhmetelashvili et all., 2007; Hartman et all., 2008; Rueda-Orozco et all., 2008).

all., 1993; Royall et all., 2002; Longstaff, 2006).

Passingham, 1993).

The pattern of basal-thalamic-cortical connections carrying out motor, emotional and cognitive functions is mutual for these functions (Alexander et all., 1996; DeLong et all., 1998; Herrero et all., 2002; Groenewegen, 2003; Saint-Cyr, 2003; Groenewegen & van Dongen, 2007; Laskowska et all., 2008), and the control of these functions consists in the choice of the sequence of behaviours adapted to a particular situational context (Heilman et

The control of motor, emotional and cognitive functions is hierarchical and the processing of information connected with these functions can be executed sequentially or simultaneously in various structures of the brain. Clinical observations and the results of the investigations carried out in persons with a damage of the basal ganglia confirm the participation of subcortical structures not only in motor but also cognitive functions (Oberg and Divac, 1979; Berns and Sejnowski, 1995; Frank et all., 2001). Also the results of computer simulations of cognitive actions such as: maintenance of information in operating memory, remembering, planning of sequences of behaviors and making decisions indicate a participation of the basal ganglia in the control of cognitive functions (Prescott et all., 2002; Jessup et all., 2011). The function of the basal ganglia in the control of motor, emotional and cognitive functions relates to cooperation with frontal cerebral cortex areas (Brown et all., 1997). The basal ganglia cooperating with the prefrontal cerebral cortex make it possible to initiate motor, emotional and cognitive actions. A complex system of stimulations (inhibition and disinhibition) limits the flow of excessive information which disables the initiation of an activity. A complex choice mechanism makes it possible to initiate a particular kind of movement without detailed determination of the whole sequence of the movements executed (Bullock & Grosberg, 1988; Hikosaka, 1989; Chevalier & Deniau, 1990;

According to the concept of mental (motor, emotional, cognitive) functions control carried out by cortico-subcortical loops a proper action of all the structures of these loops is a condition necessary for their efficient functioning (Brown et all., 1997). Mental processes, such as for example: making decisions, the choice of motor action, the change of behavior, working memory can be disturbed in a similar degree, independently from which structure of a particular loop is damaged (ibid.). Because of a specific structure of striatal medium spiny neurons (having a very large quantity of synaptic connections) and synaptic processes like long-term synaptic potentiation (LTP) and long-lasting synaptic depression (LTD), similar to those carried out in the cerebral cortex, the striatum (including the ventral striatum) is functionally the main structure of cortico-subcortical loops (Picconi et all., 2005) that controls motor, emotional and cognitive functions. Due to numerous inter-striatal and cortico-striatal connections and described above plastic mechanisms enabling enhancement or reduction of stimulations, the striatum together with the cerebral cortex controls the functions of particular cortico-subcortical loops, substantially influencing mental processes (Brown et all., 1997; Prescott et all., 2002).

The results obtained from the experiments on a model of shoulder movement simulation indicate that the higher dopamine concentration is in the striatum the shorter is the time necessary to initiate a movement (ibid.). This means that the time necessary to initiate a movement becomes shorter simultaneously with an increase of the striatum stimulation (ibid.). Moreover, the simulation of a decrease in dopamine concentration in the striatum in the same model causes bradykinesia and akinesia, similar to that observed in the Parkinson's disease (Prescott et all., 2002). The key function of the striatum is to produce signals which reach, through the thalamus, the cerebral cortex (ibid.). It was suggested that the choice of a particular pattern of action takes place in the striatum due to so called gating mechanism. Activation of the neurons responsible for processing of a particular pattern leads to inhibition of other striatal neurons. The mechanism makes it possible e.g. to process efficiently information introduced to the working memory with simultaneous inhibition of the inflow of new information before completing a presently executed task (Frank et all., 2001). If the gate is closed, new information does not influence the memory essentially and that is why it allows a stable maintenance of information being processed (ibid.). Opening of the gate enables data updating (ibid.). The inhibition of new information inflow prevents their interference with previously gathered data (ibid.). This mechanism enables continuous selection of the processed information optimizing in this way execution of mental actions (ibid.). The disturbance of the mechanism of selective actualization of data in proper time (both too frequent and too rare data updating) increases the number of errors made in various tasks. This disorder relates to initiation of both movement and thought processes. A confirmation of the concept equity is psychic akinesia observed in persons with a damage of the striatum (Brown and Marsden, 1990). However, inhibition of the internal part of the globus pallidus leads to disinhibition of frontal loops activity, which results in continuous updating of data. Continuous updating of data is caused by the lack of blocking in access to the working memory (Frank et all., 2001). Results of these disorders can be Absentmindedness, impulsivity and hyperactivity (occurings in the disorders such as: Huntington's disease, Tourette syndrome, attention deficit hyperactivity disorder) can be the results of the above mentioned disorders (ibid.). Diminishing of dopamine concentration in the striatum causes that the gate regulating the access to the working memory is opened and closed in the wrong time, and in permanent inhibition of the internal part of the globus pallidus it is opened all the time (ibid.).

The gating model does not fully explain the selection of data, which arises from the fact that dopamine is secreted in large areas of the prefrontal cortex. In this model, different areas of the prefrontal cortex are activated simultaneously, making it impossible to select one

Functional Anatomy, Physiology and Clinical Aspects of Basal Ganglia 101

According to the described above hypothesis on action selection, an alternative explanation for the functional organization of the basal ganglia was proposed. Instead of traditionally selected pathways (*direct* and *indirect)* the existence of neuronal circuits responsible for the choice (called selection circuit) and control (control circuit) of the activities undertaken is postulated. The selection circuit (traditionally called: direct pathway), leading from D1 dopamine receptors of striatal medium spiny neurons to the entopeduncular nucleus (EP) (or internal globus pallidus in primates) and a reticular part of the substantia nigra pars reticularis (SNR), receiving also the stimulation from the subthalamic nucleus reaching the EP and SNr, creates a selection mechanism acting as the feedforward selection circuit, which allows, on the basis of information coming into the system, the choice of motor or cognitive schema, before it starts to be executed. In this perspective, the neural connections of the striatum with the globus pallidus (Gp) and the nucleus subthalamicus (traditionally called indirect pathway) are included in the loop responsible for the choice control (control circuit) (Gurney et all., 1998, Prescott et all., 2002). Based on the analysis of the results of the experiments simulating the described above way of the striatum function regulation, two functions of the control loop were distinguished: 1) inhibition of STN by the GP using negative feedback allows to regulate the number of impulses outgoing from STN through the channels of information flow (associated with the activity of striatal medium spiny cells) (ibid.); 2) inhibition of EP/SNr by the GP as a part of a selection support mechanism. The increase in concentration of dopamine in the striatum facilitates the choice of information channels, which will be disinhibited, while the decrease in concentration of dopamine hinders the choice (ibid.). The reason of errors occurring in the selection of behaviors adjusted to a particular situation or difficulties in completing already chosen behaviors in the Parkinson's disease, in which concentration of dopamine in the nigrostriatal pathway is decreased, are tried to be explained with the above presented model (Prescott et all., 2002).

Akhmetelashvili O.K., Melkadze I.A., Davitashvili M.T. & Oniani T.I. Effects of electric

Alexander G. E., DeLong M. & Strick P. L. Parallel organization of functionally segregated circuits linking basal ganglia and Cortex *Ann. Rev. Neurosci.* 1986, 9, 357-381. Albin R. L., Young A. B. & Penney J. B. The functional anatomy of basal ganglia disorders.

Alexander G.E., Crutcher M.D. & DeLong M.R. Basal ganglia-thalamocortical circuits:

Babinski J. Sur le réflexe cutané plantaire dans certaines affections du systeme nerveux central. *Comptes rendus des Séances et Mémoires de la Société de Biologie* 1896, 207-208. Beiser D.G., Houk J.C. Model of cortical-basal ganglionic processing: encoding the serial order of sensory events. *The Journal of Neurophysiology,* 1998, 79 (6), 3168-3188. Berns G.S,. Sejnowski T.J. How the basal ganglia make decisions. W: Damasio A., Damasio

Bochenek A., Reicher M. *Anatomia człowieka*. Wyd. lek. PZWL, Warsaw 2006.

stimulation and destruction of caudate nucleus on short-term memory. *Bull Exp.* 

parallel substrates for motor, oculomotor, "prefrontal" and "limbic" functions.

H., Christen Y., red. *The Neurobiology of Decision Making*. Berlin, Springer-Verlag,

**6. References** 

*Biol. Med.* 2007, 143 (3): 302-304.

*Trends. Neurosci.* 1989, 12, 366-375.

1995, 101-113.

*Progress in Brain Research,* 1990, 85, 119-46.

particular piece of information (ibid.). This mechanism does not meet the functional requirements of selective updating of data, which is, as explained above, a simultaneous updating of certain information and maintaining the remaining data unaltered (ibid.).

The gating model is complemented by the concept of selective gating mechanism based on the functioning of parallel basal-thalamic-cortical loops (Frank et all., 2001). Inhibitory nature of the connections leading from the striatum to the internal part of the globus pallidus (GPi), the reticular part of the substantia nigra (SNr) and the thalamus has a decisive influence on the processes of gating. For this reason, the activity of neurons in the striatum causes the stimulation of neurons in the thalamus (via dual inhibition) (Deniau & Chevalier, 1985, Chevalier & Deniau, 1990). Disinhibition of the thalamus evokes gating function that enables emergence of other functions, although it does not induce them directly.

Selective gating mechanism explains the role of the basal ganglia in the initiation of the process of collecting new information in the working memory and the possibility of its inflow of new information before completing a presently executed task (Frank et all., 2001). If the fast update (Frank et all., 2001). In the absence of stimulation of striatal neurons the gate remains closed and the frontal cerebral cortex maintains the information currently being processed. This is possible due to existence of multiple parallel loops (ibid.).

In a classical division five parallel basal-thalamic-cortical loops were proposed (Alexander, 1986). However, on the basis of anatomical characteristics, existence of many subloops inside this five basic circuits was assumed. Due to this a relatively precise control of the working memory is possible (Beiser & Houk, 1998). It was assumed that control of the working memory depends on maintaining constant stimulation of neurons of the prefrontal cortex (Frank et al, 2001), which is the result of a continuous self-stimulation via a feedback loop (ibid.). Maintaining of this state is possible due to active feedback connections between the neurons of the frontal cerebral cortex and the thalamus. (ibid.).

The importance of the basal ganglia in senso-motor processes consists in the fact that they decide on the choice of a particular activity (Prescott et all., 2002). In the computational model, which was based on the observation of two different, competing with each other behaviors, it is assumed that coded is only one of them (i.e. the one whose execution by the motor system is being currently more important) (Redgrave et all., 1999). At the cellular level, a complex choice mechanism that allows solving a conflict between competing behaviors through fast and decisive switch between pre-selected actions, may be related to the degree of polarization of the neuron cell membrane. Resting membrane potential of striatal spiny neurons fluctuate from the values close to depolarization state so-called "up" state to hyperpolarization - "down" state (Calabresi et all., 2007). The selection mechanism consists of four steps: 1) selection of the cells in "up" state and exclusion of the cells in "down" state; 2) local inhibition within the striatum, which selectively increases the probability of stimulation of some spiny neurons and decreases the probability of stimulation of the other ones (accordingly to the cognitive model it increases the probability of the flow of information through specific channels (perceived as groups of spiny cells) (Redgrave et all., 1999); 3) localized inhibition induced by striatal medium spiny neurons (D1) together with dispersed stimulating impulses, incoming from the subthalamic nucleus, which act like a feed- forward loop, controlling the information coming out of the basal ganglia (from the internal globus pallidus and the substantia nigra reticularis); 4) local mutual inhibition occurring in the output basal ganglia (thalamus), which additionally restricts the selection criteria (ibid.).

According to the described above hypothesis on action selection, an alternative explanation for the functional organization of the basal ganglia was proposed. Instead of traditionally selected pathways (*direct* and *indirect)* the existence of neuronal circuits responsible for the choice (called selection circuit) and control (control circuit) of the activities undertaken is postulated. The selection circuit (traditionally called: direct pathway), leading from D1 dopamine receptors of striatal medium spiny neurons to the entopeduncular nucleus (EP) (or internal globus pallidus in primates) and a reticular part of the substantia nigra pars reticularis (SNR), receiving also the stimulation from the subthalamic nucleus reaching the EP and SNr, creates a selection mechanism acting as the feedforward selection circuit, which allows, on the basis of information coming into the system, the choice of motor or cognitive schema, before it starts to be executed. In this perspective, the neural connections of the striatum with the globus pallidus (Gp) and the nucleus subthalamicus (traditionally called indirect pathway) are included in the loop responsible for the choice control (control circuit) (Gurney et all., 1998, Prescott et all., 2002). Based on the analysis of the results of the experiments simulating the described above way of the striatum function regulation, two functions of the control loop were distinguished: 1) inhibition of STN by the GP using negative feedback allows to regulate the number of impulses outgoing from STN through the channels of information flow (associated with the activity of striatal medium spiny cells) (ibid.); 2) inhibition of EP/SNr by the GP as a part of a selection support mechanism. The increase in concentration of dopamine in the striatum facilitates the choice of information channels, which will be disinhibited, while the decrease in concentration of dopamine hinders the choice (ibid.). The reason of errors occurring in the selection of behaviors adjusted to a particular situation or difficulties in completing already chosen behaviors in the Parkinson's disease, in which concentration of dopamine in the nigrostriatal pathway is decreased, are tried to be explained with the above presented model (Prescott et all., 2002).

#### **6. References**

100 Neuroimaging for Clinicians – Combining Research and Practice

particular piece of information (ibid.). This mechanism does not meet the functional requirements of selective updating of data, which is, as explained above, a simultaneous updating of certain information and maintaining the remaining data unaltered (ibid.). The gating model is complemented by the concept of selective gating mechanism based on the functioning of parallel basal-thalamic-cortical loops (Frank et all., 2001). Inhibitory nature of the connections leading from the striatum to the internal part of the globus pallidus (GPi), the reticular part of the substantia nigra (SNr) and the thalamus has a decisive influence on the processes of gating. For this reason, the activity of neurons in the striatum causes the stimulation of neurons in the thalamus (via dual inhibition) (Deniau & Chevalier, 1985, Chevalier & Deniau, 1990). Disinhibition of the thalamus evokes gating function that enables emergence of other functions, although it does not induce them

Selective gating mechanism explains the role of the basal ganglia in the initiation of the process of collecting new information in the working memory and the possibility of its inflow of new information before completing a presently executed task (Frank et all., 2001). If the fast update (Frank et all., 2001). In the absence of stimulation of striatal neurons the gate remains closed and the frontal cerebral cortex maintains the information currently

In a classical division five parallel basal-thalamic-cortical loops were proposed (Alexander, 1986). However, on the basis of anatomical characteristics, existence of many subloops inside this five basic circuits was assumed. Due to this a relatively precise control of the working memory is possible (Beiser & Houk, 1998). It was assumed that control of the working memory depends on maintaining constant stimulation of neurons of the prefrontal cortex (Frank et al, 2001), which is the result of a continuous self-stimulation via a feedback loop (ibid.). Maintaining of this state is possible due to active feedback connections between

The importance of the basal ganglia in senso-motor processes consists in the fact that they decide on the choice of a particular activity (Prescott et all., 2002). In the computational model, which was based on the observation of two different, competing with each other behaviors, it is assumed that coded is only one of them (i.e. the one whose execution by the motor system is being currently more important) (Redgrave et all., 1999). At the cellular level, a complex choice mechanism that allows solving a conflict between competing behaviors through fast and decisive switch between pre-selected actions, may be related to the degree of polarization of the neuron cell membrane. Resting membrane potential of striatal spiny neurons fluctuate from the values close to depolarization state so-called "up" state to hyperpolarization - "down" state (Calabresi et all., 2007). The selection mechanism consists of four steps: 1) selection of the cells in "up" state and exclusion of the cells in "down" state; 2) local inhibition within the striatum, which selectively increases the probability of stimulation of some spiny neurons and decreases the probability of stimulation of the other ones (accordingly to the cognitive model it increases the probability of the flow of information through specific channels (perceived as groups of spiny cells) (Redgrave et all., 1999); 3) localized inhibition induced by striatal medium spiny neurons (D1) together with dispersed stimulating impulses, incoming from the subthalamic nucleus, which act like a feed- forward loop, controlling the information coming out of the basal ganglia (from the internal globus pallidus and the substantia nigra reticularis); 4) local mutual inhibition occurring in the output basal ganglia (thalamus), which additionally

being processed. This is possible due to existence of multiple parallel loops (ibid.).

the neurons of the frontal cerebral cortex and the thalamus. (ibid.).

restricts the selection criteria (ibid.).

directly.


Functional Anatomy, Physiology and Clinical Aspects of Basal Ganglia 103

Groenewegen H.J. The ventral striatum as an interface between the limbic and motor

Groenewegen H.J., van Dongen Y.C. *Role of the basal ganglia*. W: W. van Laar (red.) *Parkinsonism and related disorders*. VU University Press Amsterdam 2007, 21-54.

Haber S. N. The primate basal ganglia: parallel and integrative networks. *J. Chem.* 

Haber S.N., Calzavara R. The cortico-basal ganglia integrative network The role of the

Hartley A. A., Speer N. K. Locating and fractionating working memory using functional

Hartman R.E., Rojas H., Tang J. & Zhang J. Long-term behavioral characterization of a rat model of intracerebral hemorrhage. *Acta. Neurochir. Suppl*. 2008, 105, 125-126. Heilman K.M., Bowers D. & Valenstein E. *Emotional disorders associated with neurological* 

Herrero M. T., Barcia C. & Navarro J. M. Functional anatomy of thalamus and basal ganglia.

Hikosaka O. Role of basal ganglia in intiation of voluntary movements. W: Arbib M.A.,

Hikosaka O., Takikawa Y. & Kawagoe R. Role of the basal ganglia in the control of purposive saccadic eye movements. *Physiological Revievs,* 2000, 80 (3), 953-978. Jessup R.K., O'Doherty J.P. Human Dorsal Striatal Activity during Choice Discriminates

Joel D., Weiner I. The organization of the basal ganglia-thalamocortical circuits: Open interconnected rather than closed segregated. *Neuroscience.* 1994, 63, 363-79. Kostowski W. Zaburzenia procesów neuroprzekaźnictwa w chorobie Parkinsona. In: Friedman A., red. *Choroba Parkinsona.*, Bielsko-Biała, α-Medica Press, 1999, 18. Kowalska D.M., Kuśmierek P. *Anatomiczne Podstawy Pamięci.* In: T. Górska, A. Grabowska, J.

Kozubski W., Paweł P. & Liberski P.P. (edit.) *Neurologia. Podręcznik dla studentów Medycyny*.

Laskowska I., Ciesielski M. Gorzelańczyk E.J. Udział jąder podstawy w regulacji funkcji

Laskowska I., Gorzelańczyk E.J. Rola jąder podstawy w regulacji funkcji poznawczych.

L'Hermitte F. 'Utilization behaviour' and its relation to lesions of the frontal lobes. *Brain.*

L'Hermitte F. Human autonomy and the frontal lobes, part II: patient behavior in complex

and social situations: the environmental dependency syndrome. *Ann. Neurol.* 1986,

emocjonalnych. *Neuropsych. Neuropsychol.* 2008, 3 (3-4), 107-115.

Warsaw, Wydaw. Lekarskie PZWL. 2006, 84-85.

*Neuropsych. Neuropsychol.* 2009, 4, 1, 26-35. *LeDoux* J.E. *The Emotional Brain.* Simon & Schuster, New York 1996.

1983, 106 (Pt 2), 237-255.

19, 335–343.

neuroimaging: storage, maintenance and executive functions. *Microsc. Res. Tech.* 

*disease*. In: K.M Heilman, E. Valenstein (edit.) *Clinical Neuropsychology* Oxford

Amari S., red*. Dynamic interaction in neural networks: models and data.* Berlin,

Reinforcement Learning Behavior from the Gambler's Fallacy. J Neurosci. 2011,

Zagrodzka (Edit.) *Mózg a zachowanie*. Wydawnictwo Naukowe PWN, Warsaw 2006,

Gurney K.N., Prescott J. & Redgrave P. The basal ganglia viewed as an action. 1998

systems. *CNS Spectr*. 2007, 12 (12): 887-892.

thalamus. *Brain Res. Bull.* 2009, 78 (2-3), 69-74.

*Neuroanat.* 2003, 26 (4), 317-330.

University Press, New York 1993.

Springer-Verlag, 1989, 153-167.

*Childs. Nerv. Syst*. 2002, 18 (8), 386-404.

2000, 51(1), 45-53.

31(17), 6296-304.

349-374.


Brown R.G., Marsden C.D. Cognitive function in Parkinson's disease: from description to

Brown L.L., Schneider J.S. & Lidsky T.I. Sensory and cognitive functions of the basal ganglia.

Bullock D., Grossberg S. Neural dynamics of planned arm movement: Emergent invariants

Bunzeck N., Doeller C.F., Dolan R.J. & Duzel E. Contextual interaction between novelty and

Calabresi P, Picconi B., Tozzi A. & Di Filippo M. Dopamine-mediated regulation of corticostriatal synaptic plasticity. *Trends in Neurosciences,* 2007, 30 (5), 211-219. Chevalier G., Deniau J.M. Disinhibition as a basic process in the expression of striatal

Cummings J. L. Anatomic and behavioral aspects of frontalsubcortical circuits. W: Grafman

DeLong M.R., Primate models of movement disorders of basal ganglia origin. Trends

DeLong M., Wichmann T. & Vitek J.L. (1998) Pathophysiological basis of neurosurgical

Deniau J.M., Chevalier G. Disinhibition as a basic process in the expression of striatal

Drewe E. A. Go no-go learning after frontal lobe lesions in humans. *Cortex.* 1975, 11, 8 –16.

Fuster J. M. Memory and planning: two temporal perspectives of frontal lobe function. In:

Frank M.J., Loughry B. & O'Reilly R.C. Interactions between frontal cortex and basal ganglia

Friedman A. Epidemiologia, etiopatogeneza, rozpoznawanie i leczenie choroby Parkinsona. In: Friedman A., red. *Choroba Parkinsona*. Bielsko-Biała, α-Medica Press, 1999. Goldman-Rakic P. S. Anatomical and functional circuits in prefrontal cortex of non-human

Gołąb B.K. *Anatomia czynnościowa ośrodkowego układu nerwowego.* 2000, Wydawnictwo

Groenewegen H. J. The basal ganglia and motor control. *Neural. Plast.* 2003, 10 (1-2), 107-120.

Annals of the New York Academy of Sciences, vol. 769, 1995, 1–13.

and speed-accuracy properties during trajectory information. *Psychological Review,* 

reward processing within the mesolimbic system. Hum Brain Mapp. 2011 Apr 21.

J., Holyoak K. J., Boller F., red. *Structure and Functions of the Human Prefrontal Cortex*.

treatment of Parkinson's disease, In: *Srereotactic and Functional Neurosurgery*, Gildenberg, Tasker, pp. (1139-1146), The McGraw-Hill Comp. Inc, New York. DeLong M.R., Wichmann T. Circuits and circuit disorders of the Basal Ganglia. *Archives of* 

functions: II. The striato-nigral influence on thalamocortical cells of ventromedial

Jasper H. H., Riggio S., Goldman-Rakic P. S., red. *Epilepsy and the Functional* 

in working memory A computational model Cognitive, Affective & Behavioral,

primates: relevance to epilepsy. In: Jasper H. H., Riggio S., Goldman-Rakic P. S., edit. *Epilepsy and the Functional Anatomy of the Frontal Lobe.* 1995a, New York, Raven,

theory. *Trends in Neurosciences,* 1990, 13, 21-29.

functions. *Trends in Neurosciences,* 1990*,* 13, 277-280.

thalamic nucleus. *Brain Research,* 1985, 334, 227-233.

Fix J.D*. Neuroanatomia*. 1997, Urban & Partner, Wrocław. Fuster J. M. *The Prefrontal Cortex.* 1980, Raven, New York.

*Neuroscience.* 2001, 1 (2), 137-160.

Lekarskie PZWL, Warsaw.

85–96.

Elliott R. Executive functions and their disorders. *Br. Med. Bull.* 2003, 65, 49-59.

*Anatomy of the Frontal Lobe.* 1995, New York, Raven, 9–18.

*Cur. Opinion. Neurobiol.* 1997, 7, 157-163.

1988*,* 95, 49-90.

doi: 10.1002/hbm.21288.

Neurosci., 1990 Jul; 13 (7), 281-5.

*Neurology,* 2007, 64 (1), 20-24.


Functional Anatomy, Physiology and Clinical Aspects of Basal Ganglia 105

Picconi B., Pisani A. , Barone I., Bonsi P., Centonze D., Bernardi G. & Calabresi P.

Prescott T.J., Gurney, K. & Redgrave, P. Basal ganglia. W: Arbib M. A., red., *The Handbook of Brain Theory and Neural Networks*, Cambridge, MA, MIT Press, 2002, 147-151. Redgrave P., Prescott T. & Gurney K. Basal ganglia: A vertebrate solution to the selection in

Rogers R. D., Own A. M., Middleton H. C., Williams E. J., Pickard J. D., Sahaklan B. J. &

Rueda-Orozco P.E., Montes-Rodriguez C.J., Soria-Gomez E., Méndez-Díaz M. & Prospéro-

Sadikot A. F., Rymar V.V. The primate centromedian-parafascicular complex anatomical

Sadowski B. *Biologiczne mechanizmy zachowania się ludzi i zwierząt.* Wydawnictwo Naukowe

Saint-Cyr J.A. Frontal-striatal circuit functions context, sequence, and consequence. *J. Int.* 

Schotland J.L., Rymer W. Z. Wipe and flexion reflexion reflexes of the frog, II. Response to

Schiffer A.M., Schubotz R.I. Caudate nucleus signals for breaches of expectation in a

Sławek J. Zabiegi stereotaktyczne w chorobie Parkinsona - zasady kwalifikacji chorych w świetle dotychczasowych badań. *Neurol. Neurochir. Pol*. 2003, 1, 215-227. Smith E. E., Jonides J. Storage and executive processes in the frontal lobes. *Science*. 1999, 283

Smith Y., Raju D.V., Pare J.F. & Sidibe M. The thalamostriatal system: a highly specific network of the basal ganglia circuitry. *Trends Neurosci.* 2004, 27(9), 520–527. Sobstyl M., Ząbek M. & Koziara H. Patofizjologiczne podstawy operacyjnego leczenia

Stahl S.M. Stahl's essentials phychopharmacology. Neuroscientific basis and practical

Szołna A. *Historia stereotaksji i neurochirurgii czynnościowej*. *Leczenie operacyjne zaburzeń ruchu*:

*choroba Parkinsona* In: M. Harat (edit.). *Neurochirurgia czynnościowa.* TOM,

choroby Parkinsona. *Neurol. Neurochir. Pol.* 2003, 37 (1), 203-213.

applications. University of California, San Diego, 2008.

movement observation paradigm. Front Hum Neurosci. 2011, 5, 38. Sławek J., Bojko E. & Szady J. Częstość występowania otępienia u chorych z chorobą

disease. *Movement Disorders,* 2005,20 (4), 395-402.

problem? *Neuroscience,* 1999, 89, 1009-1023.

*Neuropharmacolog.* 2008, 55(1), 55-62.

*Neuropsych. Soc.* 2003, 9, 103-127.

perturbation. *J. Neurophysiol.* 1993, 69, 1736-1748.

Parkinsona. *Neurol. Neurochir. Pol*. 2001, 35, 569-581.

14 (4), 377-405.

PWN, Warsaw 2001.

(5408): 1657-1661.

Bydgoszcz 2007, 9-19.

130.

Pathological synaptic plasticity in the striatum: implications for Parkinson's

Robbins T. W. Choosing between small, likely rewards and large, unlikely rewards activates inferior and orbital prefrontal cortex. *J. Neurosci.* 1999, 19, 9029– 9038. Royall D. R., Lauterbach E. C., Cummings J. L., Reeve A., Rummans T. A., Kaufer D. I.,

LaFrance W. C. & Jr, Coffey C. E. Executive control function: a review of its promise and challenges for clinical research. A report from the Committee on Research of the American Neuropsychiatric Association. *J. Neuropsychiatry Clin. Neurosci.* 2002,

García O. Impairment of endocannabinoids activity in the dorsolateral striatum delays extinction of behavior in a procedural memory task in rats.

organization with a note on neuromodulation. *Brain Res. Bull.* 2009, 78 (2-3), 122-

Longstaff A., *Krótkie wykłady. Neurobiologia*. 2003, Wydawnictwo Naukowe PWN, Warsaw.

Longstaff A., Instant Notes in Neuroscience. BIOS Scientific Publ, 2006.


Longstaff A., *Krótkie wykłady. Neurobiologia*. 2003, Wydawnictwo Naukowe PWN, Warsaw.

Mandir A. S., Lenz F. A. Clinical pathophysiology in Parkinson's Disease. In: Gildenberg

McFarland N.R., Haber S.N. Thalamic relay nuclei of the basal ganglia form both reciprocal

McNab F., Klingberg T. Prefrontal cortex and basal ganglia control access to working

Middleton F. A., Strick P. L. Anatomical evidence for cerebellar and basal ganglia involvement in higher cognitive function. *Science.* 1994, 266, 458–461. Milner B. Aspects of human frontal lobe function. W: Jasper H. H., Riggio S., Goldman-Rakic

Mink J. W. The basal ganglia: focused selection and inhibition of competing motor

Mink J.W. Basal ganglia. In: Zigmond M.J., Bloom F.E., Landis S.C., Roberts J.L., Squire L.R., red. *Fundamental Neuroscience.* San Diego, Academic Press, 1999, 951-972. Morgane P.J., Galler J.R. & Mokler D.J. A review of systems and networks of the limbic forebrain/limbic midbrain. *Progress in Neurobiology,* 2005, 75 (2), 143-160. Oberg R.G.E., Divac I. (1979). Cognitive functions of the neostriatum. In: *The Neostriatum.*

Obeso J.A., Rodriguez M.C. & DeLong M.R. Basal ganglia pathophysiology. A critical

Obeso J. A., Rodriguez- Oroz M. C., Rodriguez M. Lanciego J.L., Artieda J., Gonzalo N.

Olié J.P., Costa e Silva J.A. & Marcher J.P. (2004). Interpretacja kliniczna danych

Olzak M., Gorzelańczyk E. J. Zwoje podstawy i wzgórze a pamięć operacyjna i funkcje

Parent A., Hazrati L. Functional anatomy of the basal ganglia. I. The cortico-basal ganglia-

Peterson B. S., Skudlarski P., Gatenby J. C., Zhang H., Anderson A. W & Gore J. C. An fMRI

Petri H.L., Mishkin M. Behaviorism, cognitivism and the neuropsychology of memory. *Am.*

study of Stroop Word-Color Interference: evidence for anterior cingulate subregions subserving multiple distributed attentional systems. *Biol. Psychiatry.*

Passingham R.E. *Frontal lobes and voluntary action.* 1993, Oxford University Press, Oxford. Percheron G., Filion M. Parallel processing in the basal ganglia: up to a point. *Trends.* 

wykonawcze: przegląd badań. *Aktualn. Neurol*. 2005, 4(5), 282-289.

thalamo-cortical loop. *Brain Res. Rev.* 1995, 20, 91-127.

& Olanow C.W. Pathophysiology of the basal ganglia in Parkinson's disease. *Trends* 

biologicznych dotyczących neuroplastyczności. In: *Neuroplastyczność. Patofizjologia depresji w nowym ujęciu*. Olié, Costa e Silva, Marcher, pp. 72, Gdańsk, Wydawnictwo

Ph.L., Tasker R.R., edit. *Textbook of Stereotactic and Functional Neurosurgery.* New

and nonreciprocal cortical connections, linking multiple frontal cortical areas. *J.* 

P. S. *Epilepsy and the Functional Anatomy of the Frontal Lobe.* 1995, New York, Raven,

Longstaff A., Instant Notes in Neuroscience. BIOS Scientific Publ, 2006.

York, McGraw-Hill, 1998, 1133 – 1137.

memory. *Nature Neurosci.* 2008, 11, 103-107.

programs. *Prog. Neurobiol.* 1996, 50, 381–425.

review. *Adv. Neurol.* 1997, 74, 3- 18.

*Neurosci.* 2000b, 47(10), 8-19.

*Neurosci.* 1991, 14 (2): 55-59.

1999, 45, 1237–1258.

*Sci*. 1994, 82, 30-37.

Via Medica.

Divac, Oberg, pp. (291-312), Pergamon Press, New York.

*Neurosci*. 2002, 22(18), 8117-8132.

67–81.


**7** 

*France* 

**Cognitive Integration in the Human Primary** 

Although many different definitions of cognition exist, there is a general acceptance that cognition can be defined as a higher function with respect to both the primary stages of sensory information processing and the final stage of motor output. This idea has been the basis of many well known psychological models where one can identify "input boxes" (i.e., visual, auditory, somatosensory information), and "output boxes" (i.e., motor commands), with, intermediate, high level (attention, language, memory, …) and low-level (motor intention, preparation) cognitive functions (see, for instance, the information processing model of Smidt and Lee (2005), or the model for central representation of goal-directed

Although these models are, without doubt, well suited to the study of cognitive processes from a psychological standpoint, they are not very helpful from a neuroscientific point of view. Indeed, ever since the very first investigations into the functioning of the living brain, the main aim has been to localize cognitive functions into the cortical structures of the brain. There exist at least **two** problems related to this approach. Firstly, and this is not a recent objection (e.g., Posner and Raichle (1998) page 16), it is doubtful whether the cognitive functions as presently conceived have a meaning for the brain. Let us take for example the so-called "eye-hand coordination". This "function" is much studied today and many publications report attempts to localize it in the brain. But, for a normally developed brain, this is not a specific function which is needed at specific moments and which is necessarily implemented in a specific brain structure. All input is continuously put in relation with each other as a function of the particular output. It seems more likely that eye-hand coordination is controlled in a continuous, implicit and distributed way. It is pertinent here to mention the ecological approach of perception (Gibson, 1986). This approach is based on the concept of "affordance" that characterizes the object of perception as a whole of many possible actions and interactions, and is in rupture with the cognitive approach. Indeed, according to the latter approach, the brain organizes the perception of the world, whereas in the ecological approach, the world organizes the perception: The role of the brain is to extract the information presented by the world. This theory suggests that the traditional approach of studying cerebral functioning is not very appropriate: the cognitive functions that we define do not have much sense for the brain and, what's more, we generally put subjects in

**1. Introduction** 

movements of Jeannerod (1990)).

**Sensory and Motor Areas: An Overview** 

Jozina B. De Graaf1 and Mireille Bonnard2

*2Mediterranean Institute for Cognitive Neuroscience CNRS – Aix-Marseille University, Marseille,* 

*1Institute of Movement Sciences* 

