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Cortical Modulation of Dorsal Column Nuclei: a Computational Study

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Cortical Modulation of Dorsal Column Nuclei: a Computational Study J Comput Neurosci DOI 10.1007/s10827-006-7058-5 Cortical modulation of dorsal column nuclei: A computational study Eduardo Sanchez´ · Senen´ Barro · Jorge Marino˜ · Antonio Canedo Received: 25 June 2004 / Revised: 20 November 2005 / Accepted: 10 January 2006 / Published online: 22 April 2006 C Springer Science + Business Media, LLC 2006 Abstract We present a computational study aimed at ex- 1. Introduction ploring the sensorimotor cortex modulation of the behaviour of dorsal column nuclei, specifically the impact of synaptic The dorsal column nuclei (DCN), located in the most caudal parameters, during both sleep and waking conditions. On the area of the brainstem, are constituted by the gracile (GN) basis of the circuit proposed by Canedo et al. (2000), we have and the cuneate (CN) nuclei. Both nuclei have a similar developed realistic computational models that have been topographic organization (neuronal types and connections), tested with simultaneous electrocorticographic as well as differing in the origin of their primary afferents: GN neurons intracellular cuneate recordings performed in anaesthetized receive inputs from lower thoracic and lumbar segments, cats. The results show that, (1) under sleep conditions, the while CN inputs arise from the upper part of the trunk and model can block the transmission of afferent sensory in- forelimbs. formation and, (2) operations expected during wakefulness, The DCN constitutes a part of the dorsal column-medial such as filtering and facilitation, can be performed if synaptic lemniscus (DC-ML) pathway, which processes cutaneous parameters are appropriately tuned. and proprioceptive information. This pathway is consti- tuted by peripheral receptors, the aforementioned DCN, the Keywords Sensorimotor cortex . Cuneate nucleus . ventro-postero-lateral (VPL) nucleus of the thalamus, and Cutaneous transmission . Cortical modulation . Sleep the somatosensory cortex (SSC); connected through several rhythms bundles of ascending fibers: primary afferents, fibers form- ing the medial lemniscus (ML) (which connect the DCN with the VPL), and thalamo-cortical axons. This pathway is modulated by a large number of descending fibers from the sensorimotor cortex (SMC), contacting topographically re- Action Editor: Steve Redman lated areas of subcortical structures (Bindman and Lippold, E. Sanchez´ () · S. Barro 1981; Kuypers, 1981; Canedo, 1997). r Grupo de Sistemas Intelixentes (GSI), Departamento de SMC fibers to the VPL. Arising in layer VI of the SMC Electronica´ e computacion,´ Facultade de F´ısicas, Universidade de Santiago de Compostela, 15782 Santiago de Compostela, Spain and making synaptic contacts with thalamocortical (TC) e-mail: [email protected] neurons and local internerneurons, as well as with the r GABAergic cells of the thalamic reticular nucleus (TRN). J. Marino˜ SMC fibers to the DCN. Grouped into two main tracts Departamento de Medicina, Facultade de CC. da Saude,´ Universidade da Coruna,˜ (Canedo, 1997; Bindman and Lippold, 1981): direct cor- 15006 A Coruna,˜ Spain ticocuneate fibers and collaterals of the corticospinal tract. The former have their origin mostly in areas 1, 2 and 3b A. Canedo of primary somatosensory cortex. The latter, which do Departamento de Fisiolox´ıa, Facultade de Medicina, Universidade de Santiago de Compostela, not send collaterals to the VPL or to the TRN (Castman- 15782 Santiago de Compostela, Spain Berrevoets and Kuypers, 1976), are constituted by axons Springer J Comput Neurosci arising from area 4 (primary motor cortex (MC)), area 6 some of the classical studies, have questioned others and (premotor cortex), and areas 1, 2 and 3 (SSC) (Biedenbach have added new insights about synaptic connections and re- and De Vito, 1980). These tracts also differ in their con- lationships between the cerebral cortex and the CN. With re- duction speeds, which are slower for the corticocuneate gard to cuneate interneurons, in addition to excitatory effects, fibers (Lamas et al., 1994; Martinez et al., 1995). cortical inhibitory effects have also been observed (Canedo et al., 2000). Moreover, excitation and inhibition on the same 1.1. The cuneate nucleus and the corticocuneate interneuron have been discovered (Canedo et al., 2000)af- circuitry ter stimulating SSC and MC, respectively. Regarding CT neurons, both inhibitory (Marino˜ et al., 1999) and excita- The middle zone of the cuneate nucleus (CN), located be- tory effects (Marino˜ et al., 2000) were confirmed. How- tween the obex and 4 mm caudal to it. This area is composed ever, the mediating inhibitory mechanism is still open to de- of a core or central region, and a shell or peripheral region. bate as recent data provides evidence of postsynaptic rather The core is basically made up of clusters of cuneothala- than presynaptic inhibition (Canedo, 1997; Aguilar et al., mic (CT) neurons (Kuypers and Tuerk, 1964), also referred 2003). All these findings suggest the existence of a complex to as projection or relay neurons, which are surrounded by corticocuneate circuitry (Canedo et al., 2000), as shown in interneurons, also known as local neurons or non-thalamic Fig. 1. projection neurons. The shell, comprising the ventral and Our interest is focused on the role of corticocuneate cir- marginal regions, is mostly made up of interneurons. CT cuitry during wakefulness and sleep. It has been proposed cells project their axons to the VPL nucleus whereas in- that, during wakefulness, SMC could control the flow of terneurons exert their influence on CT neurons as well as information through the DCN by two complementary mech- other interneurons. anisms (Mountcastle and Darian-Smith, 1968; Canedo et al., The CN receives inputs from primary afferents (origi- 2000): (1) facilitation of the transmission of information nating from cutaneous and deep receptors), brainstem, and originating in those cutaneous areas that either process a SMC. Cutaneous fibers are basically confined to the central high intensity stimulus or are inside the window of cortical region of the middle cuneate nucleus, preferentially estab- attention; and (2) filtering of irrelevant information. Recent lishing axodendritic synapses (Fyffe et al., 1986). Primary findings have also shown that during sleep, descending cor- afferents carrying proprioceptive information make synaptic tical fibers play an important role by imposing oscillatory contacts within the shell. Finally, the descending direct and rhythms on the DCN (Marino˜ et al., 1996, 1999). These corticospinal collateral branches can be grouped on the ba- rhythms, which have been intensively studied in the thala- sis of their cortical origin: those originating in Brodmann’s mus (McCormick and Bal, 1997; Steriade et al., 1993), are areas 4 (MC) and 3a (proprioceptive SSC) contact the shell assumed to be in charge of blocking the ascending inputs at (Cheema et al., 1984; Martinez et al., 1995); those from area subcortical stages. 3b (cutaneous SSC) project directly to the core; and those from areas 1 and 2 (cutaneous SSC) project directly into the shell as well as into the core but within zones surrounding the cluster of CT cells. The effects of cortical descending fibers on DCN cells have previously been studied by Jabbur and Towe (1961), Towe and Jabbur (1961), and Levitt et al. (1964). These re- searchers found that SMC either excites, blocks or has no effect on the activity of DCN cells. Gordon and Jukes (1964) reported that cortical stimulation inhibits neurons in the me- dial DCN region. This inhibition was partially explained by the depolarization of primary afferent cutaneous terminals (Andersen et al., 1964a, b). Corticofugal fibers would excite DCN interneurons, which in turn would depolarize the pri- mary afferent terminals and thus decrease the synaptic effect of sensory signals. The overall effect would be presynaptic inhibition. On the other hand, it was also demonstrated that SMC could directly excite CT cells (Darian-Smith, 1966; Fig. 1 The corticocuneate circuitry. Three possible synaptic connec- tions are depicted: cortical excitation and inhibition on interneurons Winter, 1965). Recent experiments in the cuneate nucleus, (inset 1), cortical inhibition on cuneothalamic neurons (inset 2), and based on intracellular recordings in vivo (Marino˜ et al., 1999, cortical excitation on cuneothalamic neurons (inset 3). Scheme adapted 2000; Canedo et al., 2000) have confirmed the results of from Canedo et al. (2000) Springer J Comput Neurosci The aim of the present study was to analyze the behaviour 3. Primary afferents and cortical neurons. Two compart- of the circuit proposed by Canedo et al. (2000) under dif- ments for soma and axon. We took the membrane cur- ferent synaptic configurations, and particularly, to find out rent as the sum of a fast sodium current (Huguenard which synaptic parameters explain the behaviour observed et al., 1988), and a delayed rectifier potassium current in vivo in sleep as well as wakefulness. We present: (1) the (McCormick and Huguenard, 1992). computational methods used in the present study, (2) a com- putational model for the circuitry proposed by Canedo et al. 2.2. Synaptic level (2000), (3) computer experiments performed with the model, and predictions derived from them, and (4) a final discussion. To describe conductance dynamics associated with the synaptic current we have used the alpha function, which represents the conductance associated with the synaptic cur- 2. Methods rent and determines the shape of the postsynaptic potential generated after appropriate stimulation (Jack
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