Telencephalon

Largest, most rostral part of human brain. Developed from forebrain (one of three primary vesicles)
in the 4 th week. In the 5th week prosencephalon divides into telencephalon and diencephalon.
Telencephalon forms 3 parts; the middle unpaired part- telencephalon medium. In the fully
developped brain it is reduced to only thin plate that borders third ventricle anteriorly called lamina
terminalis. While paired part of telencephalon grows a lot in all directions and forms left and the
right hemispheres. Beside The secondary vesicles also central canal enlarges, into ventricular system.
The lamina terminalis developing from median telencephalon borders third ventricle which
communicates with lateral ventricle (horse shoe shaped space in hemispheres) via interventricular
foramen). Originally the smooth surface of hemisphere which lasts up to 3 rd month of
developement but afterwards the surface of the hemisphere enlarges forming lobes and gyri.

During the 2 nd month, hemisphere further grows to form the wall called pallium and cortex while
lateralbasally the hemisphere developes in striatum which gives rise to basal ganglia. Pallium then
differentiates into 3 parts; neopallium, paleopallium, archipallium. Part of pallium derived remains
on the lateral side as insular cortex. This cortex is overgrown by other lobes as opercula that meet in
the sulcus lateralis. While basal ganglia remains paramedian.

(The striatum, also known as the neostriatum or striate nucleus, is a subcortical part of the forebrain. It receives input from
the cerebral cortex and is the primary input to the basal ganglia system. In all primates, the striatum is divided by a white
matter tract called the internal capsule into two sectors called the caudate nucleus and theputamen. The term corpus
striatum is occasionally used to refer to the striatum combined with the pallidum, which is a collective term for the dorsal
pallidum (globus pallidus) and the ventral pallidum. The lenticular nucleusrefers to the putamen together with the globus
pallidus)

In 3 rd month of intrauterine developement, neopallium grows more than archi and paleopallium
and pushes the archipallium mostly medially and paleopallium basally, ventrally.

The cortex contains about 50 billion neurons and 500 billion glial cells. It has both laminar and
collaminar organisation. Columns form functional units so called modules that respond to stimuli like
movement at particular joint for example. However, some columns form multi modules meaning that
they receive inputs not only from cutaneous receptors but also proprioreceptive stimuli. Then they
provide the necessary identification of 3D object held in hand.

Cortex covers hemisphere- Superolateral surface, medial and inferior surfaces. Each hemisphere has
frontal, parietal, occipital, temporal lobes. Frontal lobe is seperated from parietal lobe by central
sulcus. The parietal lobe is seperated from occipital lobe by parietooccipital sulcus on medial surface
and inferiorly occipital lobe seperated from temporal lobe by preoccipital notch. Temporal lobe
seperated from frontal and parietal lobe by lateral sulcus that deepens in the something fossa which
contains the insula. From the medial aspect we can describe another lobe which frons the corpus
callosum and diencephalon, it is formed by gyrus cinguli that continues to the inferior surface to
gyrus parahippocampalis which turns medially as incus. Because this lobe surrounds centre of
telencephalon as a limb- called in latin limbus. This lobe also called limbic lobe. This derived the
limbic system.

There is important calcarine sulcus in the occipital lobe which is surrounded by primary tissular
cortex. Inferiorly we can see the olfactory bulb and tract which lies in olfactory sulcus then sulcus
hippicampi which seperates brainstem from parahippocampal gyrus and this is seperated from other
gyri by collateral sulcus. When operculum frontale, pariatela and temporal removed we can see
insula (hidden lobe) which has several short gyri anteriorly gyrus breve insulae and posteriorly gyrus
longi insulae. At the section through hemispheres we can see distrubiton of grey matter of
hemisphere on the surface which forms cortex and it enters all sulci at the surface. Cortex has
lamillar organisation while another grey matter inside the hemisphere and forms ncl called basal
ganglia. White matter which is between these two parts of grey matter is formed by loosely
myelinated axons that forms 3 types of pathways; 1.Projection pathways that are ascending or
descending systems because they connect cortex to lower levels of CNS. 2.Commisural pathways
connect similar sites of cortex of both hemispheres and largest commisure is corpus callosum.
3.Association pathways connect the sides of different lobes within one hemisphere.

GM: BG,cortex WM pathways: Proj, Commis,Assc

Cerebral cortex is divided developementally into the older part which develops from paleopallium
and archipallium so its paleocortex which is reduced to basal part of cortex and it is functionally
correspond to olfactory system or rhinencephalon so paleocortex forms only 1 percent of cortex.
Archicortex is mostly on the medial surface of hemisphere and then in depth of temperal lobe and
forms only about 2-3 percent of cortex. Neocortex is formed by 6 layers, archicortex and paleocortex
sometimes called together as allocortex, formed by 3-5 layers. There are also parts of transitional
cortex that are between paleocortex and neocortex it is called peripalleocortex. And between
archicortex and neocortex it is called periarchicortex. They have also 6 layers and are refered as
mesocortex.

Rhinencephalon is formed by olfactory bulb and tract and trigonum and stria olfactoria medialis and
lateralis. The archicortex of the limbic system is reduced to the induseum griseum which covers the
surface of corpus callosum and then to the hippocampal formation which is inside the temporal lobe.
Limbic lobe is already the mesocortex and neocortex that surrounds the corpus callosum and it is
formed by gyrus cinguli and gyrus parahippocampalis.
Stria olfactoria lateralis connects the olfactory triangle to the piriform lobe which is anterior
parahippocampal gyrus at uncus. All these tract and triangle represents paleocortex. Piriform lobe
forms the mesocortex but the centre of highest olfactory discrimination is in the neocortex of orbital
part of frontal lobe so in the posterior orbitofrontal cortex which where olfactory stimuli are
percieved. Olfactory stimuli is transmitted to neocortex through pirifrom lobe and mediodorsal ncl of
thalamus.

Limbic system is very wide system that place role in emotional behaviour and it develops in close
association with olfactory system. Limbic system has integrated function, it also influences the
visceral, endocrine and emotional state. hippocampal formation has also memoral function. Limbic
system includes not only structures that are located in telencephalon but also subcortical structures,
sub parts of thalamus, hypothalamus and midbrain so we talk about about limbic forebrain if we
describe structures that belong telencephalon, diencephalon. On the other hand there is limbic
midbrain located in the mesencephalon and there are very numerous connections between these
two parts.

Component of limbic forebrain is hippocampal formation which consists of hippocampus (projects on

inferior horn of lateral ventricle) and it is also called cornu ammonis after resemblence with
energyption dating with erems head. Other parts of archicortical hippocampal formation is gyrus
dentatus and subiculum which is the medial surface of parahippocampal gyrus so all these structures
are formed by archicortex of limbic forebrain. And they have role for memory. The hippocampal
formation receives inputs from the entorino cortex (periarchicortex- mesocortex of the
parahippocampal gyrus- its inferior surface. This entorino cortex receives afferentation from
association areas of neocortex. It transmits impulses to hippocampal formation through the white
matter which covers ventricular surface of hippocampus called alveolus for consolidation and then
the impulses again are transmitted to association areas of cortex mostly via fornix which is the main
pathway of limbic forebrain. It means that the hippocampus is important for consolidation of short
term memory, if hippocampal formation is injured it results in amnesia. While long term memory is
stored in association areas of neocortex.

Fornix is pathway which starts from hippocampal formation as fimbria hippocampi. Then it continues
as crus fornicis, posteriorly and superiorly both crura fornicis are connected by commisura fornicis
below splenium of corpus callosum. Crura fornicis joined to form corpus cornicis and corpus again
diverge into columnae fornicis, columnae fornicis has free part which borders anteriorly
interventricular foramen and hidden part which end in the mamillary body of hypothalamus. But
some fibers also direct to anterior commisure so called precommisural fibers and tothe septal area of
the medial surface of hemisphere.

According to classic conception the main circuit of limbic system is papetz circut which connects
gyrus parahippocampalis with mamillary body via fornix. Then mamillary body contains neurons that
send their axons to anterior nucleus of thalamus via mamillothalamic tract. This anterior ncl of
thalamus is connected to the gyrus cinguli and gyrus cinguli then sends axons to gyrus
parahippocampalis and again to the hippocampus. But it is only one circuit formed by pathways of
limbic system because limbic system is very complex system which is connected with many cortical
and subcortical areas/centres so some authers thinks papetz circuit as main pathway of limbic
system. So the papetz circuit has no specific function. According to the descent reception we speak
about Limbic forebrain which includes especially basomedial telencephalon, structures of
diencephalon; these are connected to mesencephalon which present limbic midbrain. These
structures are very important for emotion and motivation of our behavior. These structures
especially are gyrus cinguli, parahippocampalis, hippocampus, insular cortex, neocortical regions of
forebrain especially orbitofrontal cortex and basal frontotemporal regions, from ncl whose that are
involved functionally in the limbic forebrain are septal area, amygladar ncl, ventral striatum
(pallidum), nuclear complex, anterior and dorso medial ncl of thalamus, habenula, hypothalamus
especially mamillary body (ncl mamillaris).

Papetz circuit: So hippocampus sends axons via fornix to the mamillary body, there they synapse
with other neurons send send their axons to anterior ncl of thalamus, there are other neurons that
send fibres to gyrus cinguli. And gyrus cinguli is connected to parahippocampal gyrus and it is
connected again to hippocampus.

There is imp septal area which is represented by paraterminal gyrus on the medial surface of
hemisphere. Paramedial gyrus has no laminar organisation because there are septal ncl. These septal
ncl receive inputs from amygdalar complex through diagonal bend of Broca ( The diagonal band of
Broca is one of the forebrain nuclei that are derived from the ventral telencephalon during development. This structure forms
the medial margin of the anterior perforated substance.),
also receive inputs from olfactory system via stria
olfactoria medialis. Also from hippocampus via fornix and other inputs comes from midbrain from
monoaminergic neurons of midbrain via medial forebrain bundle.

The main efferent pathway of the septal area which also extends to septum pellucidum is the fornix,
through which these efferent axons reach hippocampus and another efferent pathway is stria
medullaris thalami through which these neurons reach the habenular ncl and then interpeduncular
ncl of midbrain. There are cholinergic neurons in the septal are and together with cholinergic ncl of
reticular formation they are involved in sleep awake cycles. And another ncl which also contains
cholinergic neurons and which is important for maintainance state of awaking of cortex is the basal
ncl of Mynert. (Nucleus basalis of Meynert, abbreviated NBM and also known as the nucleus basalis, is a group of
neurons in the substantia innominata of the basal forebrain which has wide projections to the neocortex and is rich
in acetylcholine and choline acetyltransferase). Both
areas are affected during Alzeimer disease which is
characterized by loss of cholinergic neurons in basal ncl of Mynert and septal areas .

Broadman areas

The neocortex forms nearly 95 percent of the each hemispehere, it may be divided according many
criteria like amount of neurons in layers of cortex according to amount of glial cells, according to
neurotransmitters or according to number of afferent and efferent fibres. So this criteria is used to
create the map and most used map is Broadmans divided neocortex into 11 regions and 52 areas acc
to cytoarchitectural differences.

You should know primary areas. Primary motor area located in the precentral gyrus, a 4 and it is
connected to secondary motor area which is rostral to this primary motor area. When primary motor
area is injured it results in paralysis of contralateral muscles.

Primary sensory are is limited to postcentral gyrus a 3,1,2. It should be

said in this order because a 3 is oriented into central sulcus received inputs from rapidly adapting
receptors, a 1 on the top of postcentral gyrus receives inputs from slowly adopting receptors, a2
receives inputs from articulations there is also area 3a which is in the floor of central sulcus receives
inputs from muscle spindles. When this primary sensory area is injured it results in loss of sensitivity-
anestezia. Secondary sensory area is behind in the parietal lobe a 5,7

In the lower part of postcentral gyrus is a 43 which is primary taste area. The both primary motor
and primary sensory area is somatotopically organised when representation of contralateral body
parts is inverted, the face and hand have relatively large representation so we call this somatotopic
organisiton as homunculus somatosensory and homunculus motor.

Its is important to know the primary visual area which is around calcarine sulcus in the occipital lobe
a17 which closely surround this sulcus and secondary visual areas are more distant from calcarine
sulcus a18,19. When primary visual area is injured it results in cortical blindness, patient cant
recognise visual stimuli but reflexes are reserved. In the primary visual area there is retinotopic
organisation meaning that one point of retina corresponds to one point in this primary visual cortex.
In the secondary visual are there is importance of perception of some complex, visual inputs like
colour or depthness.

Primary auditory area is in the superior part of temporal gyrus. Gyrus temporalis superior a41,42 so
called the transverse temporal gyri or Heschl's gyri. Again this primary auditory area is tonotopically
organised according to pitch of noise (sounds of different frequency are processed in the brain),
when its injured it causes deafness. It is surrounded by secondary auditory area, a 22 and this area
together with a 39,40 form so called sensory cortical area for speech meaning that spoken words are
understood in these regions of cortex. When this area of speech is injured, patient cannot
understand what is said -receptive aphasia. Motor area for speech localised in a 44, 45 in the inferior
frontal gyrus and both are in dominant hemisphere, when this motor centre for speech is injured,
patient cannot express himself, both by spoken or written words called expressive aphasia or
expressive agraphia. Both centres of speech are interconnected by association pathway which forms
arcuate fasciculus which is located in the depth of supramarginal gyrus. All other regions of cortex
except for primary areas forms secondary and association areas of cortex.

Association areas are responsible for higher function of cortex like decision making and abstract
thinking for eg prefrontal cortex which is located in frontal lobe rostral to motor areas is responsible
for social behaviour for abstract thinking. When prefrontal cortex suffers lesion the subject loses his
sense of social responsibility and concentration and abstraction.

Other part of telencephalon is formed by basal ganglia which is the grey matter inside the
hemisphere. Parts: Ncl caudatus (horse shoe shaped)- its head projecting into anterior horn of lateral
ventricle, body in the middle part and tail in the posterior and inferior horns. Other basal ganglion
are Putamen and Globus Pallidus which can be divided into external and internal segments
claustrum and amygdalar complex are important basal ganglia. Basal ganglia located in hemisphere
but functionally the basal ganglia are also connected to these ncl or have the same origin like
thalamus, substantia nigra, ncl subthalamicus and tegmental area of midbrain.

Globus pallidus and putamen form topographical ncl which is called ncl lentiformis. Similar to cortex
also basal ganglia develop phylonegetically from different structures. The newest basal ganglia are
caudate and putamen forms so called dorsal striatum (from neostratum).

Ncl accumbens (a region in the basal forebrain rostral to the preoptic area of the hypothalamus) located basal to this
dorsal striatum so it forms ventral striatum. Phylonegetically older is globus pallidus called pale body
because of greater amount of myelinated axons in this ncl so its lighter than putamen or caudatus. It
is Phylonegetically its older because its developped from paleostriatum. It has lateral and medial
segments or internal and external segments. It forms dorsal pallidum and ventrally located ventral
pallidum which is connected with subthalamic ncl of thalamus.

Basal ganglion which is in both limbic system is corpus amygdeloideum or amygdalar complex
because its composed of several ncl, it develops from archistriatum and connected with archicortex.
At frontal section we can see that caudate located superiorly, it projects into the anterior horn of
lateral ventricle but its of horse shoe shape. Caudate is seperated from putamen by anterior crus of
internal capsule. Globus Pallidus sepertad from Putamen by thin layer of WM- lamina medullaris
externa and lamina medullaris interna is another thin layer of WM separates the external segment
from internal one of Globus Pallidus. Another basal ganglion claustrum is seperated from putamen
by external capsule and from cortex of insula is separated by extreme capsule. Functionally also the
subthalamic ncl and substantia nigra are involved.

The basal ganglia are subcortical ncl that play primary role in the integration of somatic motor
activity. They are involved in regulation of muscle tone and posture. And play role in the initiation,
control and cessation of automatic movements. Basal ganglia involved in limbic system they
participate also in behavioural responses related to emotions. The main circuit of basal ganglia is this
cortex sends impulses to the striatum which is superior to pallidum and then impulses runs through
the thalamus motor ncl of the thalamus back to the cortex and this circuit plans movements that are
created in the 2ndary motor area before execution of movements which is performed by 1mary
motor cortex. Primary motor cortex sends commands directly to the motor ncl of spinal cord or
cranial nerves.

The amygdalar complex is located anterior to the inferior horn of lateral ventricle. So we can see the
amygdalar complex when parasaggital section through hemisphere is made. This amygdalar complex
is connected to other structures of the limbic system and it is the centre of identificaton of danger so
its fundamental for self preservation also activity of this complex connected with fear and even
sexual responses but also with memory. When its stimulated electronically animals react with
aggression. Again at the section we can see the caudate which is connected to putamen by these
fibres which are called stria that pass through the anterior limb of internal capsule. Medial to
putamen is located pallidum and topographically pallidum plus putamen forms lentiform ncl. Basally
from putamen is located ventral striatum which is ncl accumbens. Basal to pallidum is located ventral
pallidum and basal to ventral pallidum is located the basal ncl Mynert which is within the range of
substantia perforata anterior.

The white matter of telencephalon forms the region between the cortex and basal ganglia and when
the hemisphere is cut above the corpus callosum it forms this middle part called centrum semi ovale.
(The white matter, located in each hemisphere between the cerebral cortex and nuclei, as a whole has a semioval shape. It
consists of cortical projection fibers, association fibers and cortical fibers. It continues ventrally as the corona radiata)
We can distinguish commisure, projection and association fibres. The commisural fibres connect
similar areas of cortex of both hemispheres and the largest commisure is corpus callosum. We can
describe its anterior part which starts with the narrow part- rostrum then genu, trunk and the
posterior part called splenium. The fibers direct from the corpus callosum to all parts of the cortex in
all directions so anteriorly they diverge as forceps minor that their connection is with frontal lobe
and posteriorly major forceps that contain connection of occipital lobes. These fibers form the roof
of posterior horn of lateral ventricle which is called tapetum. Another commisure is commisura
fornicis between both crura fornicis which is the commisure of the especially hippocampal formation
then there is also anterior commisure which contains phylogenitically different fibres, its commisure
of both archicortex and neocortex

Projection fibres are ascending or descending so they either enter or leave the cortex. There can be
short fibres that are identified by connection between cortex or basal ganglia or reciprocal
connections between cortex and thalamus. Most important are projection fibres that connect lower
levels of CNS to the cortex that include these efference like corticospinal tract, corticonuclear which
is functionally its same as corticospinal with a difference that corticonuclear tract ends on the motor
ncl of cranial nerves. Corticoreticular tr ends in reticular formation of brainstem, corticotecto tr in the
tectum of midbrain, corticorubra in the red ncl of midbrain, corticobulbar tr which is feedback or
sensory pathways ends on the bulbar ncl of the medulla and corticopontine pathways end on the ncl
pontis and in a component of corticopontocerebellar pathway which is imp for cerebellar control of
intendent movements.

Most of these projection fibres are involved in the internal capsule which is very important white
matter of the telencephalon because it contains practically all the projection pathways for the cortex
both ascending and descending. If there is bleeding in the region of internal capsule, it has
consequences like paralysis of the all half of the body.

Internal capsule has 3 parts. Crus anterior, genu and crus posterior. The components of each parts:
anterior limb has ant thamalocortical pathway which is ascending tract and frontopontine tract
descenting from the frontal cortex to the pontine ncl. Genu contains corticonuclear tract, it is direct
motor pathway to motor ncl of cranial nerves and all other tracts are involved in posterior crus.