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Rom J Leg Med [21] 137-148 [2013] DOI: 10.4323/rjlm.2013.137 © 2013 Romanian Society of Legal Medicine

Connection interfaces between neuronal elements and structures inside greater . Evaluation in forensic psycho-affective pathology

Gheorghe S. Dragoi1, Petru Razvan Melinte2, Liviu Radu3

______Abstract: The authors achieved a macroanatomic analysis on the location and relations of neuronal structures and elements inside transitional mesocortex and in order to visualize the connection interfaces of greater limbic system. The analysis was performed on human encephalon using subsystems generally homologated by neuroanatomists: lobus limbicus, , , lobus insularis and subcortical structures. Equally, they performed a research of the literature on the implication of connection interfaces from paralimbic, limbic and archicortex areas, into forensic psycho-affective ortology and pathology. The study draws the attention to time and space development of terminology and homologation of some new concepts bound to multifunctional subsystems such as: medial memory system, prefrontal cortex and limbic midbrain area. Key Words: greater limbic system, transitional mesocortex, archicortex

euroanatomy registered remarkable progress (proneocortical or paralimbic zone and periarchicortical or Nby the diversity of morph-functional and limbic zone); hippocampal formation (with two regions: anatomic-clinical research methods towards the knowledge retro-hippocampal area that has periarchicortical structures of neuronal elements and structures implicated in bio- and hippocampal area with archicortical structures); psycho-social processes of individual self-preservation prefrontal cortex; isle lobe, medial temporal lobe memory through nutritional and psychic activities on one hand and system and subcortical structures that include: amygdale, species self-preservation through maternal and procreation hypothalamus, anterior thalamic nuclei, accumbens septi activities on the other hand. Integrating neuronal elements nucleus and septum nuclei (Diagram 1). and structures in functional subsystems has gone a long MacLean (1990) [7] develops the concept of path from “great ” paradigm (Broca, 1878) limbic system by including some subcortical structures: [1] to that of “limbic system” (MacLean, 1954) [2] and amygdale, septum, midline thalamic nuclei, habenule and “greater limbic system” (Nieuwenhuys et al., 2008) [3] hypothalamus. The integration of connection interfaces and last but not least to description of limbic neuronal between transitional mesocortex and archicortical elements circuits (Papez, 1937 [4]; Nauta, 1958, [5], 1979 [6]). and structures, was possible after naming new anatomic- Difficulties about the belonging of neuronal clinical entities under the term of medial temporal cortex structures to limbic subsystems cumbered the knowledge (Yukie, 2000) [8]. and terminology of connection interfaces. In this respect He gathers numerous structures that belong one imposed the restructuring of neuronal subsystems to: paralimbic or proneocortical area (perirhinal area into multifunctional complexes: transitional mesocortex (A35, 36), posterior para-hippocampal cortex); limbic 1) Romanian Academy of Medical Sciences * Corresponding author: Prof. MD, PhD. e-mail :[email protected] 2) University of Medicine and Pharmacy of Craiova, Departement of Anatomy 3) Emergency Hospital , Departement of Legal Medicine, Slatina, Olt, Romania 137 Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system

TRANSITIONAL MESOCORTEX

PRONEOCORTICAL PERIARCHICORTICAL ARCHICORTICAL ZONE ZONE ZONE SUBSISTEMS

CINGULATE PCC ACC vACC A24

vPCC A23 PCG RSC A29;30 PC A35;36 LOBUS LIMBICUS TH TF PHG SC PRESUBICULUM HIPPOCAMPAL FORMATION ENTORHINAL AREA A28 DENTATUS GYRUS HR AMMON’S HORN PLC A 32 PREFRONTAL ILC A25 EXTENSII CORTEX

IG FC IC A 13 LOBUS INSULARIS HYPOTHALAMUS SUBCORTICAL STRUCTURES THALAMUS

ACCUMBENS SEPTI

SEPTUM

Diagram 1. Greater Limbic System. PCC = Posterior CingulateDiagrame Cortex; vPCC no 1. =Greater ventral LimbicPosterior System ; ACC = Anterior Cingulate Cortex; vACC = ventral Anterior Cingulate Cortex; PCG = Posterior Cingulate Cortex; RSC = Retro Splenial Cortex; PC = ; PHG = ; SC = Subiculum Complex; HR = Region; IG = ; FC = Fasciola Cinerea; PLC = Perilimbic Cortex; ILC = ; ICPCC = Insular = Posterior Cortex; Cingulate TH = Temporal Cortex; area vPCC medialis; = ventral TF =Posterior Temporal Cingulate area lateralis. Cortex; ACC = Anterior Cingulate Cortex; vACC = ventral Anterior Cingulate Cortex; PCG = Posterior Cingulate Cortex; or periarchicorticalRSC area = Retro (subiculum Splenial Cortex; , entorhinal PC = Perirhinal cortex Cortex;macroanatomic PHG = Parahippocampal knowledge Gyrus;of mesocortical SC = (paralimbic (A28), parasubiculum,Subiculum presubiculum); Complex; HR archicortical = Hippocampus area Region; and IG =limbic) Indusium and Griseum; archicortical FC = Fasciola structures Cinerea; and their (, AmmonPLC = Perilimbic horn) and Cortex; subcortical ILC = Infralimbicstructures: Cortex; integration IC = Insular in anatomic-functional Cortex; TH = Temporal systems. area amygdale complex.medialis; TF = Temporal area lateralis. Nauta (1979) [6] promoted an extension of limbic MATERIAL AND METHODS system concept by adding a continuous neuronal string named “limbic axis”, to “limbic telencephalon arch” The study was carried out in human biologic (hippocampal formation and amygdale complex without material respecting research deontology criteria. We cingulate and para-hippocampal gyrus); it contains, analyzed macroscopically and mesocopically 16 adult from rostral to caudal: septum region, preoptic region, encephalon (40-60 years old), 8 fetus encephalon and hypothalamus and several mesencephalon para-median 6 new-born encephalon, previously fixed in 5% saline structures: mesencephalic central grey and dorsal raphe formaldehyde solution. Visualization of neuronal structures nucleus (Nauta – “limbic midbrain area”). and subsystems integrated in greater limbic system was Through those extensions, Nauta lays the possible using the following methods: macroscopic and foundation of a great functional system called “limbic sculptural dissection, 3D sectioning and dissection after system – midbrain circuit”. freezing at -18ºC (Klingler’s method, 1935) [9]. The purpose of the paper is to draw the attention Macroanatomic imagery was achieved by towards neuronal structures that can offer information Canon digital camera EOS Mark II equipped with macro regarding the evaluation of normal and pathologic psycho- ultrasonic lens EF 100 mm, F/2,8. Image processing was affective processes. possible using Professional Digital Photo Software and The objectives of the paper were raised by the Adobe Photoshop CS4. 138 Romanian Journal of Legal Medicine Vol. XXI, No 2(2013)

RESULTS the inferior convex face forms the superior wall of hippocampi. We identified three parts of dentate gyrus: The macroanatomic study was done on the anterior (Giacomini’s band); posterior (fasciola cinerea) location of neuronal subsystems and structures that belong and middle. The anterior part forms the tail of dentate both to transitional mesocortex (proneocortical and/or gyrus. The posterior part (fasciola cinerea) is smooth, it periarchicortical) and archicortex. forms a small landmark that starts underneath corpus We analyzed the location and relations of five callosum, goes retrograde, surrounds splenium corpori neuronal subsystems anatomically and functionally callosi and then passes on the superior face of corpus integrated in greater limbic system: lobus limbicus, callosum where it forms “indusium griseum”. hippocampus, prefrontal cortex, lobus insularis and 3. Prefrontal cortex subsystem subcortical structures complex (Diagram 1). Prefrontal cortex subsystem frequently named as 1. Lobus limbicus subsystem “granular frontal cortex” projection cortex for medial dorsal When examining the medial face of cerebral thalamic nucleus (MD-projection cortex) is considered hemispheres we easily identified some of the limbic lobe nowadays “high order heteromodal association area” structures: gyrus cinguli and isthmus gyri cinguli (Fig. 1, (Mesulam, 1983) [10]. Although the term prefrontal was A, D, E, F). introduced in 1868 by Richard Owen, the prefrontal area Gyrus cinguli (synonym in French terminology was narrowed to the most anterior part of . The to “circonvolution du corp calleux”, “le lobul du corp definition of prefrontal cortex was based on cell-architecture calleux”, “lobe calleux”) circumscribes that is the presence of IV granular layer (Jacobsen, and is limited dorsally by sulcus cinguli (synonym in 1935) [11]. Prefrontal cortex is nowadays considered a French terminology to “scissure sous frontale”, “scissure projection area for medial dorsal thalamic nucleus (Rose calloso-marginale”) and ventrally by sulcus corporis and Wollsey, 1948)[12]. Macroanatomic analysis of callosi. The anterior extremity has successive relations to frontal lobe allowed us to identify three major regions rostrum corpori callosi and genu corpori callosi (Fig. 1, A, integrated in prefrontal cortex: lateral, medial and orbital. E). The posterior extremity, after it goes around splenium 4. Lobus insularis subsystem corpori callosi, directs ventrally and it continues through The isle lobe or lobule was described in 1796 by a passage fold homologated as “isthmus gyri cinguli” Johann Christian Reil. In French literature, it is named (synonym in French terminology to “le pli temporo- either as “central lobe of Gratiolet” or as striate body limbic”) to parahippocampal gyrus (Fig. 1 A, F). “lobule of Cruveilhier”. Medial face of gyrus cinguli is sometimes crossed Macroanatomic visualization was possible after by superficial grooves with variable directions, either removing the borders of lateral fissure (Sylvius) that belong parallel to gyrus trajectory or transverse or stellate (Fig. 1 to frontal and temporal lobes. It appears surrounded by a A, D). Ventral surface of gyrus cinguli forms the roof of the circular depression called “sulcus circulare insulae”. The groove for corpus callosum where one can find the anterior tip of the isle (Limen insulae) serves as a starting point, cerebral artery; it has relations to “indusium griseum” in a fan fashion, for isle gyruses (gyrus longus insulae (synonym in French terminology to “tractus gris de and gyrus breves insulae) and in between them there are Lancisi”) that is a prolonged part of dentate gyrus (Fig. 1 D). isle grooves. One of those grooves is deeper; it appears 2. Hippocampus proprius subsystem precocious in ontogenesis and is named the central groove Visualization of hippocampus was achieved (Sulcus centralis insulae). On frontal or horizontal sections after the dissection of temporal lobe in its internal and one can notice the relations of isle lobes to and inferior parts. It appears as a prominence in the inferior (Fig. 2 and 3). horn of lateral ventricle. On serried sections it is covered 5. Subcortical structures subsystem by a layer of white substance (alveus). On the internal During the anatomy history, the following border, there is hippocampal fimbria that is fringed on subcortical structures were included in the greater limbic its inferior face. Two borders and two faces can be easily system: amygdale, hypothalamus, thalamus ,accumbens identified on transverse sections. The superior face is septi nucleus, and septum nuclei. free and contributes to border Bichat’s fissure (Fissura a. The amygdale nucleus system (Corpus transversa cerebri). The inferior face adheres in its middle amygdaloideum) described by Burdach (1819) becomes part to Ammon’s horn. The internal border is free and the visible either after opening the anterior part of the external border serves for the insertion of pia mater that inferior horn of lateral ventricle where it determines the forms choroid plexus. Fimbria ends anteriorly to the tip prominence called amygdale tubercle, or on encephalon of Ammon’s horn, precisely where joins Ammon’s frontal sections where one can notice its relations to horn, and posteriorly to (Fig. 5). hippocampus. It appears in the dorsal and medial part of Between hippocampal fimbria and Ammon’s temporal lobe where it forms the rostral-medial and rostral- horn we visualize the dentate gyrus that appears crossed dorsal parts of the inferior horn of lateral ventricle (Fig. 2). by fissures limiting transverse folds. The superior face b. Hypothalamus was integrated in the limbic of dentate gyrus is covered by hippocampal fimbria and system (Papez, 1937 [4]; MacLean, 1990 [7]). It appears as

139 Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system a subcortical structure interposed between telencephalon – determined remarkable progresses in elaborating neuronal rostral and mesencephalon – caudal. Analyzing the sagittal limbic pathways. Important contributions are attributed median sections one can notice hypothalamus location to Papez (1937) [4], Nauta (1958) [5], (1979) [6], in the most anterior part of diencephalon contributing MacLean (1954) [2] and Brodmann (1909)[14]. Equally, to the formation of the lateral wall of the . the complexity and diversity of anatomic and functional The dorsal (superior) limit is obvious by the presence of interrelations between the structures and subsystems of hypothalamic groove (Sulcus hypothalamicus). The rostral greater limbic system, imposed the elaboration of new (anterior) border of hypothalamus area coincides with the anatomic and functional complexes such as: prefrontal vertical frontal plane that passes through Monro foramen cortex complex, subicular complex, hippocampal (Foramen interventriculare) and through the middle of formation, medial temporal lobe memory system (Squire optic chiasm. Ventral limit corresponds to the inferior and Zola Morgan, 1994) [15], great limbic lobe (Broca, wall of the third ventricle and contains the optic chiasm, 1878) [1], limbic system (MacLean 1954) [2], entorhinal infundibular recess and mammillary bodies (Fig. 1). cortex system – hippocampus and transitional mesocortex. c. Thalamus takes parts in limbic circuits by “limbic Due to those progresses, there were described nuclei” that include the anterior nucleus complex and numerous connection interfaces between: proneocortical dorsal lateral nucleus interconnected to cingulate cortex, area and cortical area; proneocortical area and , presubiculum and parasubiculum. periarchicortical area; proneocortical area and archicortex; Macroanatomic analysis of thalamus location lobus limbicus and neocortex; hippocampus and neocortex and its relations was carried out on sagittal, frontal and by entorhinal cortex; sensory cortex and hippocampus by horizontal sections. Medial face of thalamus was evaluated perirhinal area. after making a middle sagittal section through encephalon. The main nodal points were stated in the limbic It is part of the lateral wall of the third ventricle. The limits system: amygdale complex, septum and anterior thalamic of medial face of thalamus are easily recognized: dorsal nucleus. (superior) by habenule, ventral (inferior) by hypothalamic groove (Fig. 1). In the anterior half it is united to medial A. Considerations on connection interfaces in face of contralateral thalamus by adhesion interthalamica. paralimbic part of greater limbic system. Implications in When examining the frontal and horizontal sections, one psycho-affective ortology and pathology can observe the relation of thalamus lateral face to internal Paralimbic structures implicated in psycho- capsule that separates thalamus from – affective processes are: ventral posterior cingulate cortex situated dorsal and lateral, and lentiform nucleus – situated (VPCC); perirhinal cortex; prelimbic cortex and insular lateral and ventral. The convex dorsal face is limited cortex ( Diagram 1 ) laterally by . 1. Ventral posterior cingulate cortex (VPCC) is d. septi was identified where present in posterior and ventral part of cingulate gyrus the caudate nucleus meets the anterior part of putamen, and corresponds to Brodmann areas 29, 30, 23 and 31. In lateral from ; it is easy to visualize 1993, B. A. Vogt [16] imposed a new division of posterior ventral and medial from the head of caudate nucleus. cingulate cortex in two sectors: posterior (PCC) limited to Heimer (1978) [13], based on structural, histochemical areas 23 and 31 and retrosplenial (RSC) that contains only and interconnection studies, considers that accumbens Brodmann areas 29 and 30. In order to avoid confusions, nucleus and as parts of corpus . B. A. Vogt named the complex of the two sectors It represents an interface between the limbic system and with the term “posterior cingulate gyrus “(PCG). The motor system. posterior sector receives afferent fibers from hippocampal e. Septum is divided into a dorsal part (septum formation. Both sectors are involved in remembrance and pellucidum) and a ventral part (septum verum). The topographic orientation. ventral part is situated rostral from 2. Perirhinal cortex (PC) is situated in the inside paraterminal gyrus; it is bordered dorsally by anterior part of parahippocampal gyrus as an interface rostrum corpori callosi, anteriorly by pre-commissure between sensory cortex and hippocampus. It is integrated part of hippocampus and caudally by in parahippocampal region of hippocampal formation and preoptic region. Ventral and lateral it is relation to together with entorhinal cortex and subicular complex. accumbens nucleus. It occupies an intermediate position It plays an important role in information remembrance between limbic system and striatum (extrapyramidal (working memory). system) and it is considered as a neuronal station between 3. Perilimbic cortex (PLC) was identified as hippocampal formation and hypothalamus. a region of prefrontal cortex (mPFC) and located in the dorsal part of cingulate gyrus (dACC) and in Brodmann DISCUSSIONS area 32. It integrates information regarding motivation and mnemonics. Description of connection interfaces between 4. (IL) is part of isle lobe (Reil, the structures and subsystems of greater limbic system 1796). By it connections to thalamus and amygdale central

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Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system

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C ED CC 2 6 EE Figure 3. Hippocampus macroanatomic relations in the inferior horn of lateral ventricle (A-C, E) and isle lobe relations to claustrum andFigure putamen no. (C, D) 3. onHippocampus horizontal sections macroanatomicthrough cerebral hemispheres. relations 1. Polus in frontalis. the inferior 2. Lobus insularis.horn of 3. FigureFigure no. no. 3. 3. HippocampusHippocampus macroanatomicmacroanatomic relationsrelations inin thethe inferiorinferior hornhorn ofof Plexuslateral choroideum ventricle of ventriculus (A-C, lateralis E) and 4. Fimbriaisle lobe hippocampi. relations 5. Claustrum. to claustrum 6. Putamen. and 7. Hippocampus putamen proprius (C, D) (Cornu on ammonis).lateral 8. ventricle Subiculum. Macrophotos(A-C, E) byand Canon isle EOS lobe Digital relations Camera Macro to claustrumUltrasonic Lens, and EF4 100 putamen mm, F/2,8. (C, D) on horizontal sections4 through cerebral hemispheres. 1. Polus frontalis. 2. Lobus horizontalhorizontal sectionssections throughthrough cerebralcerebral hemispheres.hemispheres. 1.1. PolusPolus frontalis.frontalis. 22.. LobusLobus insularis. 3. Plexus choroideum of ventriculus lateralis 4. Fimbria hippocampi. 1435. insinsularis.ularis. 3.3. PPlexuslexus choroideumchoroideum ofof ventriculusventriculus lateralislateralis 4.4. FimbriaFimbria hippocampi.hippocampi. 5.5. Claustrum. 6. Putamen. 7. Hippocampus proprius (Cornu ammonis). 8. Subiculum. Claustrum.Claustrum. 76.6. Putamen.Putamen.8 7.7. HippocampusHippocampus propriusproprius (Cornu(Cornu7 ammonis).ammonis). 8.8. Subiculum.Subiculum. Macrophotos by Canon EOS Digital Camera Macro Ultrasonic 8Lens, EF 100 mm, Macrophotos by Canon EOS Digital Camera Macro Ultrasonic Lens, EF 100 mm, F/2,8. F/2,8.

C E Figure no. 3. Hippocampus macroanatomic relations in the inferior horn of lateral ventricle (A-C, E) and isle lobe relations to claustrum and putamen (C, D) on horizontal sections through cerebral hemispheres. 1. Polus frontalis. 2. Lobus insularis. 3. Plexus choroideum of ventriculus lateralis 4. Fimbria hippocampi. 5. Claustrum. 6. Putamen. 7. Hippocampus proprius (Cornu ammonis). 8. Subiculum. Macrophotos by Canon EOS Digital Camera Macro Ultrasonic Lens, EF 100 mm, F/2,8.

Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system

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Romanian Journal of Legal Medicine Vol. XXI, No 2(2013)

1 1 1. Fimbria hippocampi 11 3 1. Fimbria hippocampihippocampi 33 1. Fimbria2. Gyrus hippocampi dentatus 2. Gyrus dentatusdentatus 2. Gyrus3. dentatusVentriculus lateralis et 6 3. Ventriculus3. Ventriculus lateralis et lateralisplexuslateralis choroideus etet 66 plexus choroideus 6 4. Cornu Ammonplexus choroideuschoroideus 4. Cornu Ammon 2 5. Sulcus4. hippocampalisCornu AmmonAmmon 22 A 5. Sulcus hippocampalis A 6. Fanta5. BichatSulcus hippocampalishippocampalis 7 4 7. Subiculum6. Fanta Bichat 77 44 6. Fanta BichatBichat 5 A 8. Gyrus7. parahippocampalisSubiculum 555 7. Subiculumum 9. Sulcus8. collateralisGyrus parahippocampalis 8. Gyrus parahippocampalisparahippocampalis 8 10. Bandeleta9. Sulcus optică collateralis 88 9. Sulcus collateraliscollateralis 11. Eminentia10. Bandeleta collateralis optică 12. Digitationis10. Bandeleta hippocampi opticăoptică 11. Eminentia collateralis 11. Eminentia collateraliscollateralis AA 9 12. Digitationis hippocampi A 9 12. Digitationis hippocampihippocampi 666 B BB

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Figure 5. Hippocampus shape and relations on successive serried sections from posterior (A) to anterior (D); location and hippocampusFigureFigureFigure relations to 5.5.5. gyrus HippocampusHippocampusHippocampus parahippocampalis, shapeshape Bichat’s andand slit, relations relationssulcus hippocampalis on successive and sulculs serried serriedcolateralis. sectionssections Macropghotos by Canonfromfromfrom T-70. posteriorposteriorposterior Lens autotelephoto (A)(A)(A) tototo F/2,8. anterioranterioranterior (Professor (D);(D); Dragoi’s locationlocation collection). and hippocampus relations toto gyrusgyrus parahippocampalis,parahippocampalis,parahippocampalis, Bichat’sBichat’sBichat’s slit,slit, sulcussulcus hippocampalishippocampalis and sulculs colateralis.colateralis. MacropMacropMacropghotosghotosghotos bybyby CanonCanonCanon TTT---70.70.70. LensLens autotelephotoautotelephoto F/2,8. (Professor Dragoi’sDragoi’s 145 collection).collection).collection).

Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system

Ventriculus Ventriculus lateralis Cornu Ammonis lateralis Cornu Ammonis 1 Fimbria Fimbria 2 hippocampi 2 hippocampi CA3 CA3 3 4 CA2 CA2 CA4 CA4

5 5 6 CA1 7 CA1 7 Sulcus Sulcus hippocampalis Gyrus dentatus hippocampalis Gyrus dentatus A A Ventriculus Ventriculus Alveus lateralis Alveus lateralis 1 2 Fimbria Fimbria CA3 hippocampi CA2 CA3 3 hippocampi CA2 3

CA4 CA4 CA1 CA1

Subiculum 5 Subiculum 5 6 Sulcus Presubiculum Aria 6 7 Sulcus Presubiculum Aria 7 hippocampalis Parasubiculum entorhinalis hippocampalis B Parasubiculum entorhinalis B Figure 6. FigureMicroanatomic no. architecture 6. Microanatomic of hippocampus proprius. architecture CA1 – CA of4 = hippocampusRegio hippocampi proprius; proprius. Strata CA1hippocampi: – 1. StratumFigure oriens. 2 Stratumno. 6. pyramidale. Microanatomic 3 Stratum molecular. architecture 4. Hilus. Strata of hippocampusgyri dentate: 5. Stratum proprius. multiforme. CA1 6. Stratum – CA 4 = Regio hippocampi proprius; Strata hippocampi : 1. Stratum oriens. 2 granulare.CA 4 7. = Stratum Regio molecular. hippocampi Crezil violet stain.proprius; Strata hippocampi : 1. Stratum oriens. 2 ImagesStratum acquired pyramidale by Nikon Eclipse . 6003 microscope.Stratum 2Dmolecular. reconstruction. 4Oc.. 7,Hilus. Ob. 4 (B), Ob.Strata 10 (A). gyrix 28 (B); dentate x 70 (A) (Professor: 5. Stratum pyramidale . 3 Stratum molecular. 4. Hilus. Strata gyri dentate: 5. Dragoi’sStratum collection). multiforme. 6. Stratum granulare. 7. Stratum molecular. Crezil violet stain. Stratum multiforme. 6. Stratum granulare. 7. Stratum molecular. Crezil violet stain. Images acquired by Nikon Eclipse 600 microscope. 2D reconstruction. Oc. Images acquired by Nikon Eclipse 600 microscope. 2D reconstruction. Oc. 7, Ob. 4 (B), Ob. 10 (A). x 28 (B); x 70 (A) (Professor Dragoi’s collection). 1467, Ob. 4 (B), Ob. 10 (A). x 28 (B); x 70 (A) (Professor Dragoi’s collection).

Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system nucleus, it intervenes in behavior psychic activities to memory center and the gateway to hippocampus. express feelings. It is important due to its roles in creating In primates it is located in the terminal, rostral body representation and emotional subjective experience. part of temporal lobe. From topographic point of view it From anatomic and functional point of the island has two is divided in two regions: medial and lateral. The absence parts: a granular posterior part that contains somatic and of neuronal layer IV is the microanatomic characteristic of sensorial areas and an anterior non granular part with entorhinal cortex. The afferent fibers come from association premotor areas involved in speech. In pathology, the isle cortical areas: perirhinal cortex, parahippocampal cortex lobe is implicated in progressive non-fluent aphasia and in and prefrontal cortex as well as hippocampus. The efferent drug addiction (cocaine, opioid substances). fibers from layers II and III project into dentate gyrus and hippocampus proprius. Layer II projects into CA B. Observations on connection interfaces in 3 region of hippocampus proprius. In addition, layer III limbic area of greater limbic system projects into subiculum and CA 1 region of hippocampus Neuronal structures from limbic area of proprius. Entorhinal system – hippocampus proprius plays transitional mesocortex are: anterior cingulate cortex an essential role in memory and its optimization during (ACC), retrosplenial cingulate cortex (RSC), subicular sleep. Entorhinal cortex is an important interface between complex, entorhinal cortex, infralimbic cortex and hippocampus proprius and neocortex. subcortical structures (amygdale complex, hypothalamus, 5. Infralimbic cortex (area 25) belongs to ventral thalamus, accumbens septi nucleus and septum). and medial prefrontal cortex. It inhibits emotional responses 1. Anterior cingulate cortex (ACC) is divided into acting on subcortical structures (Ivan Vidal Gonzales et a dorsal sector (dACC) that corresponds to al. 2006)[11]. It was proven that electric stimulation of 32 and a ventral sector (vACC) that belongs to Brodmann infralimbic cortex, reduces fear and consolidates memory area 24. It is involved in cognitive functions through (Quick, Mueller, 2007)[21]. dorsal sector and in emotional functions through ventral 6. Subcortical structures intervene in expression sector. Lesions to anterior cingulate cortex produce: of feelings and emotions, in aggressive behavior, in mnesic apathy, loss of appetite, akinetic mutism and/or vegetative consolidation processes, in transfer and processing of derangements. sensory peripheral information and/or in defense behavior. 2. Retrosplenial cingulate cortex (RSC) is a In pathology, the subcortical structures are involved in structural part of posterior cingulate gyrus together with Wernicke-Korsakoff syndrome (mammillary bodies) and posterior cingulate cortex (PCC) (B. A. Vogt, 1990)[17]. It Kluver-Bucy syndrome (amygdale). correspond to Brodmann areas 29 and 30 and it is involved in retrograde amnesia. C. The determinism of the implication of 3. Subicular complex includes three important subcortical structures connection interfaces structures: subiculum, presubiculum and parasubiculum. The connection between archicortical structures a. Subiculum is a transition area between neocortex to greater limbic system is possible due to the presence in (6 neuronal layers) and archicortex (3 neuronal layers). It ontogenesis and phylogenetic of fiber fascicles that bound plays an important role both in normal conditions (working hippocampus to other parts of encephalon, including: memory and space relations) and in pathology (epilepsy). fornix, mamillaris princeps fascicle and cingulum. It is the basal part of hippocampal formation situated Fornix is a compact fascicle of nervous fibers that between entorhinal cortex and Ammon’s horn. The afferent connect hippocampus to hypothalamus. Posteriorly from fibers come from CA1 and entorhinal cortex (layer III) and interventricular foramen a considerable amount of fibers the efferent fibers go to hippocampus proprius. leave columna fornicis and reach the anterior nucleus of b. Presubiculum is part of parahippocampal thalamus and “bed nucleus of stria terminale (BST)”. Some area and has a role in processing information regarding fibers cross hypothalamus to reach mammillary bodies. space orientation. The efferent fibers project into medial The efferent fibers from mammillary bodies region of entorhinal cortex (in rat) and into hippocampal form a compact fascicle named as “fasciculus mamillaris formation. princeps”. It passes dorsally and after a short distance it c. Parasubiculum is the major component splits in: mammillary-thalamic tract and mammillary- of subicular complex. It is a transition area between tegmental tract. subiculum and entorhinal area (Zilles, 1990) [18]. The first one reaches the anterior thalamic nucleus Electrophysiological proofs suggest similarities to and it is part of Papez neuronal circuit. The second curves entorhinal cortex in rat (Funahashi and Steward, 1997)[5]. caudally and enters tegmentum to dorsal tegmental It receives numerous cortical and subcortical impulses and nucleus and pons reticular tegmentum nucleus described sends major projections to superficial layer of entorhinal by Bechterev. cortex (Amaral and Witter, 1995)[19]. It is involved in Cingulum is a fascicle of association fibers that topographic orientation, space location and integration of becomes visible only after a minute sculptural dissection. sense information (Chrobak and Buzsakie, 1994) [20]. It crosses through gyrus cinguli and parahippocampal 4. Entorhinal cortex (Area 28) is an important gyrus to reach septum area and uncus of temporal lobe.

147 Dragoi G.S. et al Connection interfaces between neuronal elements and structures inside greater limbic system

CONCLUSIONS or paralimbic and periarchicortical or limbic on one hand and on the other hand, it allowed the understanding of 1. The knowledge of connection interfaces bio-psycho-social processes of species and individual self- between the neuronal elements and structures of greater preservation. limbic system is important for the understanding of the 4. The concept of limbic axis elaborated by complexity in time and space of psycho-affective processes Nanta (1979) as attached to telencephalon limbic arch, whose evaluation supposes great responsibility. determined the statement of a great functional system 2. Minimizing and maximizing connection named limbic system midbrain circuit that afterwards was interfaces between the structures of greater limbic system integrated in greater limbic system. are imposed by the stereodistribution and elements 5. The anatomic and functional unity of the assembly in anatomic and functional neuronal subsystems. structures forming greater limbic system is possible due to 3. The concept of “transitional mesocortex” the existence of numerous connection interfaces between allowed the enrolment of the structures and elements of elements, structures and subsystems with variable greater limbic system in two great areas: proneocortical topographic locations.

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