J. Exp. Zool. India Vol. 25, No. 2, pp. 2113-2127, 2022 www.connectjournals.

com/jez ISSN 0972-0030

DocID: https://connectjournals.com/03895.2022.25.2113 eISSN 0976-1780

AGE AFFECTS THE NEURONAL SOMA DIAMETER IN CEREBRAL

CORTEX OF 15 AND 30-DAY-OLD CHICK, GALLUS GALLUS DOMESTICUS
Adarsh Kumar, Kavita Tamta, Hemlata Arya and Ram Chandra Maurya*
Neurobiology Laboratory, Department of Zoology , Kumaun University,Soban Singh Jeena Campus, Almora, India.
*e-mail : mauryarc@gmail.com
(Received 10 March 2022, Revised 4 May 2022, Accepted 12 May 2022)

ABSTRACT : The present study examines the affect of age-factor on neuronal soma diameter in different regions of the
cerebral cortex of chick, Gallus gallus domesticus, by using Cresyl-Violet-staining technique. A total of ten male chicks (5
fifteen and 5 thirty-day-old) were sacrificed in the present study. The brains were processed with Cresyl-Violet-staining
method. The anatomical analysis reveals that the cerebrum is composed of two symmetrically right and left cerebral hemispheres.
The chick cerebral hemisphere has a cavity inside called as lateral ventricle whose roof is termed as pallium that includes the
external cortical areas namely the hyperpallium apicale (HA), the hippocampal formation (HF) and the corticoid complex (CC).
All the areas show three types of neuronal cells: the multipolar, the pyramidal, and the stellate neurons. All the external cortical
areas of the telencephalon show the variation in the form of non-significant as well as significant (P<0.05, P<0.01 and P<0.001)
increase or decrease in soma diameter of the projection (pyramidal and multipolar) and stellate neurons of 30-day-old chick in
comparison to 15-day-old chick. The present study provides information that the soma diameter of cerebral cortex neurons is
larger in 15-day-old chick (early age) in comparison to soma diameter of cerebral cortex neurons of 30-day-old chick. The
present study will support the comparative, developmental and stress studies on the grounds of morphological, anatomical and
histological analysis.
Key words : Age-factor, cerebral cortex, cytoarchitecture, Nissl staining, Soma diameter.

How to cite : Adarsh Kumar, Kavita Tamta, Hemlata Arya and Ram Chandra Maurya (2022) Age affects the neuronal soma
diameter in cerebral cortex of 15 and 30-day-old chick, Gallus gallus domesticus. J. Exp. Zool. India 25, 2113-2127. DocID:
https://connectjournals.com/03895.2022.25.2113

INTRODUCTION vertebrate’s brain is composed of three regions fore, mid

In avian brain research, the domestic chicks were
used as a model for decades in the study of the
neurobiology of learning and memory (Nakamori et al,
2013). The brain is the most important part of nervous
system which has always fascinated the scientific world
toward itself and lead to new findings and information
due to which it has always been point of interest in this
field (Srivastava and Maurya, 2010). The brain possesses
remarkable ability to undergo functionally relevant
adaptations following external and/or internal stimuli
(Chattarji et al, 2015; Drevets, 2004; McEwen, 2007).
The role of central nervous system in birds such as to
integrate sensory impulses from the environment, to store
learned information and coordinate voluntary and
involuntary functions and movements, are similar to those
of mammals (Koushafar and Mohammadpour, 2019). The
and hind brain. The largest part of forebrain is cerebral
hemisphere that developed as an evagination from the
anterior end of the neural tube which represent
telencephalon. The birds and mammals possess the large
brain compared to other vertebrates as per the molecular,
anatomical, and chemical studies, which might have
evolved as an adaptation to cope with novel or altered
conditions (Al-bakri, 2016).
Birds have a three-part brain, composed of olfaction,
vision, and hearing segments whose relative proportion
varies with bird’s ecology (Koushafar and
Mohammadpour, 2019). The floor of the telencephalon
is called as sub pallium and the roof is called as the
cerebral cortex (pallium). The cerebral cortex includes
internal and external corticoid areas (Al-Nakeeb and
Jasim, 2018). The external corticoid areas consist of three
2114 Adarsh Kumar et al
complexes: hyperpallium apicale, hippocampal complex
and corticoid complex (Al-bakri, 2016).
The avian hyperpallium apicale has been found to be
associated with the learning processes, integration of
information from both eyes, orientation, and in
categorizing food from non-food (Budzynski et al, 2002;
Deng and Rogers, 2000; Heyers et al, 2007; Keary and
Bischof, 2012; Maekawa et al, 2006; Watanabe et al,
2011). The functional properties of hyperpallium apicale
were very similar to that of the mammalian visual cortex,
containing orientation selective binocular neurons which
were organized in a retinotopic manner (Bischof et al,
2016). Several studies conducted in birds revealed that
the hippocampal complex of birds have been reported to
be involved in the process of learning, memory, behaviour
regulation, cognition, and navigation (Bingman et al, 2005;
Bingman and Able, 2002; El-Falougy and Benuska, 2006;
Jacobs, 2003; Sherry, 2005; Shettleworth, 2003; Singh et
al, 2014; Singh et al, 2015; Smulders, 2006, 2017;
Srivastava et al, 2007, 2014; Tömböl et al, 2000; Vargas
et al, 2004).
Most studies based on neuronal types, neuronal
density and morphology have suggested the functional
aspect of the corticoid complex of different birds like
Pigeon (Srivastava et al, 2014), Strawberry finch, Estrilda
amandava (Srivastava et al, 2014), Indian house crow,
Corvus splendens (Srivastava et al, 2014), Zebra finch,
Taeniopygia guttata (Atoji and Wild, 2005; Colombo et
al, 2001; Montagnese et al, 1996; Singh et al, 2019;
Srivastava et al, 2014; Srivastava, Maurya et al, 2009)
and recommended that the avian dorsolateral forebrain
has participated in spatial memory. The present study
represents the neuronal soma diameter in all the areas of
cerebral cortex in chick, Gallus gallus domesticus. In
addition, the present study aims to evaluate the effect of
age factor on the soma diameter of different neuronal
cells of various regions of cerebral cortex in 15-day-old
and 30-day-old male chick, Gallus gallus domesticus.
MATERIALS AND METHODS
Experimental animal model
In present research study, a total of ten male chicks,
Gallus gallus domesticus (5 fifteen and 5 thirty-day-
old) were sacrificed. They were purchased from Pahari
Poultry House (A governmental department) Hawalbag
Almora, Uttarakhand India. On the 13th and 28th day of
their hatching, the chicks were brought in two different
cages of poultry house to the laboratory and housed in
the same cages (with the same lighting and feeding
schedules as in the poultry house). The chicks were
housed under constant laboratory conditions such as room
temperature 30°C, 12:12 light and dark cycle for next
48-hours prior to the experiment to release their stress
due to transport. The chicks were sacrificed by
administering a lethal dose of ketamine (Thermo Fisher
Scientific, India), their heads were decapitated, the intact
brain was immediately removed from the skull, and
processed with Cresyl-Violet-Staining protocol. The
present research work was approved by Institutional
Animal Ethics Committee (IAEC) with Protocol No.
KUDOPS/106. All the experiments were carried out
strictly in accordance with the animal care guidelines of
the Animal Ethics Committee of Kumaun University,
Nainital, Uttarakhand, India.
Cresyl-Violet (Nissl Stain) Method
The brain was post-fixed in 10% formalin for 24-
hours at 4°C. After 24-hours of fixation, the brain was
rinsed briefly with double-distilled water (2-3 minutes),
subjected to dehydration in 30%, 50%, 70%, 90% and
100% ethanol (15 minutes in each), cleared in xylene (20
minutes), and transferred to a mixture of xylene and molten
paraffin wax (30 minutes). Thereafter, the brain was
transferred to paraffin Wax-1, Wax-2, and Wax-3 for a
duration of 6-hours (1 hour, 2 hours and 3 hours,
respectively) at 58°C and finally the brain was embedded
in paraffin wax (m.p. 58°C – 60°C). The hardened
trimmed paraffin wax block was sectioned at thickness
of 10 µm with the aid of rotary microtome and the sections
were placed on the surface of the water (warmed at 37
°C) present over Mayer’s Albumen Glycerol coated glass
slides for spreading. After complete spreading and drying
the sections were deparaffinized in xylene (15 minutes)
followed by rehydration with 100%, 90%, 70%, 50% and
30% alcohol (10 minutes in each) and washed in double-
distilled water (10 minutes). Then the sections were
stained with 0.1% Cresyl-Violet-solution (Sigma Aldrich,
USA) for 2-3 minutes at 37 °C, rinsed briefly with double-
distilled water (5 minutes). The sections were dehydrated
with 30%, 50%, 70%, 90% and 100% alcohol (5 minutes
in each grade), cleared in xylene (5 minutes), mounted
on microscopic glass slides using D.P.X (dibutylphthalate
polystyrene xylene) mounting medium, coversliped, dried,
and stored in dark at room temperature (RT) until
microscopic examination.
Microscopic analysis
Microscopic inspection of Cresyl-Violet-stained
sections was done under bright field in a computer aided
microscope (Leica DMIL Led, Germany) at 40x, 100x
and 400x primary magnifications. In all the telencephalic
external cortical areas three classes of neuronal cells
namely: stellate neurons, multipolar and pyramidal
 Age affects the neuronal soma diameter in cerebral cortex of 15 and 30-day-old chick
projections neurons were observed on the basis of their
soma shape. The microphotographs of the entire Cresyl-
Violet-stained sections were taken with the help of a
microscope (Leica stereo-zoom-S9i) aided with computer
and the boundaries of the whole sections were traced
for the drawing of stacked serial sections. Scales for each
camera lucida drawing were made by using the stage
micrometer with the help of microscope before the
drawings were entered into the computer. All the drawings
were scanned by using a scanner and their background
was cleared with the help of Adobe Photoshop 7.0
computer software. All of the scaled microphotographs
and their drawings were used for the formation of
photoplates, which is labelled with the help of Adobe
Photoshop 7.0 computer software.
Neurohistological data measurement
For histological data measurement, we have chosen
well stained neuronal soma at random in all the regions
under study and analyzed fifty well stained neuronal soma
for each type of neuron in 15 as well as in 30-day-old
chick. Soma diameter for each neuron was measured by
using Leica Application Suite-X (LAS-X) computer
software inside computer-aided-microscope (Leica DMIL
Led, Germany) at 400x magnification.
Statistical data analysis
For statistical analysis of the neurohistological data
parametric unpaired t-test with welch’s correction was
applied to find out the differences in neuronal soma
diameter of 15 and 30-day-old chick. A minimum criterion
of probability P<0.05, P<0.01 and P<0.001, was accepted
as an indicative of significant difference. All the results
were shown as the Mean ± Standard Error of the Mean
(SEM). All the statistical analysis was performed using
Microsoft Excel, and Graph Pad Prism software’s.
RESULTS
Morphological analysis
The anatomical analysis in the present research study
revealed that the chick brain is relatively a large sized,
more or less triangular or pear shaped, broad rounded,
soft and whitish organ (Fig. 1A-D). The chick brain
consists of three major subdivisions: the rhombencephalon
(consists of metencephalon, myelencephalon and pons
varolii), the mesencephalon (consists of optic bigemina
and crura cerebri) and the prosencephalon (consists of
rhinencephalon, telencephalon or cerebral hemisphere
and diencephalon) (Fig. 1C). Telencephalon of forebrain
is one pair of large sized, pear or obtuse triangle shaped
pyriform lobes that are collectively referred to as cerebrum
which is the largest part of the brain in the presently
studied bird, chick. The cerebrum is one of the most
2115
important shapes determining factor for the entire brain
and is composed of two symmetrically right and left
cerebral hemispheres. Caudally, both the cerebral
hemispheres progressively became much broader than
its tapered rostral (anterior) end and are firmly opposed
along a median sulcus called interhemispheric
(longitudinal) fissure, while from cerebellum they are
separated by a small transverse (medial) fissure (Fig.
1A, C)
Cytoarchitectonic analysis
The complete analysis of the Cresyl-Violet-stained
telencephalic sections revealed that the chick cerebral
hemisphere has a cavity inside called lateral ventricle
(VL) or paracoel. Its thin and smooth roof is called
pallium which has a poorly developed cerebral cortex in
birds and includes the external cortical areas namely: the
rostrodorsal (wulst), the dorsomedial (hippocampal
formation) and the dorsolateral forebrain (corticoid
complex) which are situated at the tip of the telencephalic
hemisphere.
The rostrodorsal forebrain is a slight enlargement or
laminated bulge that occupies the rostro-dorsal surface
of the cerebral hemisphere extending from the midline to
lateral region of the brain and is also termed as wulst.
Externally, the lateral curved margin of the wulst (sagittal
eminence) located on either side of the inter-hemispheric
(longitudinal) fissure is demarcated from the rest of the
hemispheric surface by an indistinct groove (vallecula).
The sagittal eminence became flattened and continued
to the caudal (posterior) pole of the telencephalic
hemisphere (Fig. 1C). Functionally, the wulst has been
divided into two subfields: a rostral small somatosensory
motor area and a caudal large posterolateral visual area
termed as visual wulst (Fig. 2A, 4-5). Visual wulst is
further consisting of the four morphologically
distinguished, dorso-ventrally arranged laminae: the
hyperpallium apicale (HA), the interstitial nucleus of the
hyperpallium apicale (IHA), the hyperpallium
intercalatum (HI), and the hyperpallium densocellulare
(HD) (Fig. 2A). Hyperpallium apicale is the most
superficial lamina of the visual wulst which lies medially
at the rostral level and dorsally at the caudal level (Fig.
2A). Hyperpallium apicale is bordered by a narrower,
cellular layer called the interstitial nucleus of the
hyperpallium apicale, which lies ventro-laterally at the
rostral level and dorso-laterally at the intermediate level
(Fig. 2A). Hyperpallium intercalatum is bounded dorsally
by interstitial nucleus of the hyperpallium apicale and
ventrally by hyperpallium densocellulare (Fig. 2A). It lies
ventro-laterally at rostral level and dorso-laterally at
intermediate level. Hyperpallium densocellulare laminae
2116 Adarsh Kumar et al

Fig. 1 : Photographs showing the brain of chick, Gallus gallus domesticus: (A) dorsal view of 15-day-old chick brain showing: 1-cerebral
hemisphere, 2-optic lobes, 3-cerebellum & 4-medulla oblongata. Arrow a & b show longitudinal & transverse fissures, respectively.
(C) dorsal view of 30-day-old chick brain showing: P-prosencephalon, M-mesencephalon, R-rhombencephalon, V- vallecula & W-
wulst. (R) rostral end, (C) caudal end, (L) lateral side & (M) medial surface. (B) & (D) ventral view of 15 & 30-day-old chick brain
showing: 1-olfactory bulb, 2-cerebral hemisphere, 3-optic chiasma, 4-optic lobe, 5-cerebullum & 6-medulla oblongata.

lie ventro-laterally at rostral level and dorso-laterally at
intermediate level (Fig. 2A). Towards the caudal pole,
the interstitial nucleus of the hyperpallium apicale, the
hyperpallium intercalatum, and the hyperpallium
densocellulare disappear thus only the hyperpallium
apicale remains ventrally limited by mesopallium (Fig. 2B;
5).
The dorsomedial forebrain is relatively a narrow,
curved strip of tissue that lies on the dorso-medial surface
of the cerebral hemisphere and is separated from the
rest part of the telencephalic hemisphere by a lateral
ventricle and is also known as hippocampal complex
(HCC) (Fig. 2B; 4J-N; 5A-D). At the most rostral level,
the hippocampal complex is occupied by a small area
that is the area parahippocampalis (APH) limited
dorsolaterally by the most superficial layer of the visual
wulst that is the hyperpallium apicale (Fig. 2B; 4J-N; 5A-
D). The hippocampal complex is relatively a simple
structure in chick as clearly defined hippocampal layers
are not readily apparent in the chick hippocampus proper
(Hp) in the present study (Fig. 3C-D). At the intermediate
level, the size of the hippocampal complex is increased
dorsally and laterally from rostral towards caudal
direction. The hippocampal complex comprises two major
 Age affects the neuronal soma diameter in cerebral cortex of 15 and 30-day-old chick 2117

Fig. 2 : Microphotographs (A-D) showing the cresyl violet stained sections of chick, Gallus gallus domesticus forebrain from rostral to
caudal pole. (HA) hyperpallium apicale, (HV) hyperstriatum ventral, (LH) lamina hyperstriatica, (HP) hippocampus, (APH)
parahippocampal area, (CI) intermediate corticoid area, (L) layer, (CDL) dorsolateral corticoid area & (VL) lateral ventricle. Section
(A) is 1480 µm posterior from the first rostral section of the cerebral hemisphere, while the section (B), (C) & (D) are 2720 µm, 2600
µm & 800 µm posterior from the previous sections. Magnification 40x. Scale bar = 500 µm.

divisions: the ventrally situated hippocampus proper and
the dorsally situated area parahippocampalis (Fig. 2C;
3C-D; 5B-I). A clear boundary between the two
structures is not present in presently studied bird. A
ventrally situated V-shaped structure appeared in the
ventral two-third part of the hippocampal complex that
extends from medial to lateral ventricle is termed as
hippocampus proper (Fig. 2C; 3C; 5B-I). The
hippocampus is widest dorsally at the junction with the
area parahippocampalis and tapers ventrally towards the
septum (Fig. 2C; 3C-D; 5B-I). The dorsomedial curvature
of the telencephalon above the angle of the lateral
ventricle is known as the area parahippocampalis. The
dorsally situated area parahippocampalis over the
hippocampus proper appears to be a transitional zone
between the hippocampus proper and the adjacent
dorsolateral forebrain (Fig. 2C; 3D; 5E-I).
Parahippocampal area is only the subfield of the
hippocampal formation that is present in the rostral
hippocampal complex (Fig. 2B; 4J-N; 5A-D). At the
caudal most level of telencephalic hemisphere, the
intermediate corticoid subfield (CI) of the corticoid
complex along with the different subfields of the
hippocampal complex progressively disappeared. Thus,
caudally at the level of cerebellum, only the
parahippocampal area of the hippocampal complex
observed limited laterally by the dorsolateral corticoid
subfield (CDL) of the corticoid complex (Fig. 5L-M).
The dorsolateral forebrain or corticoid complex of
the chick is formed at the dorsolateral surface of the
telencephalic hemisphere after the progressive
replacement of the laterally situated most superficial layer
of the posteriolateral forebrain that is the hyperpallium
apicale (Fig. 2C-D; 3E-F; 5E-M). The dorsolateral
forebrain further consists of two subfields: an intermediate
corticoid subfield (CI) and a dorsolateral corticoid subfield
(CDL). The intermediate corticoid subfield is a medial
wide area that constitutes a transition zone between the
dorsolateral corticoid area of the corticoid complex and
the area parahippocampalis of the hippocampal complex
(Fig. 2C; 3E; 5E-K). In the intermediate corticoid subfield,
based on the neuronal soma size and density, three
dorsolaterally arranged cellular layers have been
recognized: Layer-I, Layer-II and Layer-III. The
outermost layer that is present towards the dorsal surface
of the cerebral hemisphere is referred to as Layer-I. The
2118 Adarsh Kumar et al

Fig. 3 : Microphotographs (A-F) showing the cresyl violet stained sections of chick, Gallus gallus domesticus forebrain from rostral to
caudal pole. (VW) visual wulst, (Hp) hippocampus proper, (APH) parahippocampal area, (CI) intermediate corticoid area, (L) layer,
(CDL) dorsolateral corticoid area & (VL) lateral ventricle. Section (A) is 1480 µm posterior from the first rostral section of the
cerebral hemisphere, while the section (B) is 2720 µm posterior from section (A), section (C) & (D) are 2600 µm posterior from
section (B), while section (E) & (F) are 800 µm posterior from the section (C) & (D). Magnification 100x. Scale bar = 100 µm.

middle Layer-II is present in the center of the intermediate extends widely on the dorsolateral surface of the caudal
corticoid subfield of the dorsolateral forebrain. The telencephalon and is separated by a lateral ventricle from
innermost layer that is present beneath the Layer-II, the underlying nidopallium until its caudal pole (Fig. 2C-
towards the ventricle is called Layer-III (Fig. 2C; 3E). D; 3F; 5H-M).
At the caudal most level of the telencephalic hemisphere, Neurohistological analysis of Telencephalon
the intermediate corticoid subfield along with the different
In presently studied bird, the Cresyl-Violet-stained
subfields of the hippocampal complex progressively
sections show the presence of three neuronal types the
disappeared. Thus, caudally at the level of cerebellum,
multipolar, the pyramidal, and the stellate neuron based
only the parahippocampal area of the hippocampal
on soma shape in all the external cortical areas of
complex observed limited laterally by the dorsolateral
telencephalon (hyperpallium apicale, hippocampus, area
corticoid subfield of the corticoid complex (Fig. 5L-M).
parahippocampalis, intermediate corticoid area, and
The dorsolateral corticoid area in chick is a thin, narrow,
dorsolateral corticoid area). The multipolar neuronal cells
superficial strip-like structure of the caudal telencephalic
possessed oval, spherical, rectangular, multiangular or
pallium adjoining the medially situated hippocampal
irregularly shaped somata (Fig. 6). The pyramidal
formation (HF). The dorsolateral corticoid subfield
 Age affects the neuronal soma diameter in cerebral cortex of 15 and 30-day-old chick 2119

Fig. 4 : Drawing of stacked serial cresyl violet stained forebrain sections from rostral to caudal pole (A-N) of 15-day-old chick, Gallus gallus
domesticus, showing the position of the (VW) visual wulst & the (APH) area parahippocampalis of the dorsomedial forebrain.
Below each section the distance from the previous section is given from rostral to caudal end. The section (A) is 760 µm posterior
from the first rostral section of the cerebral hemisphere. Scale bar = 2mm.

projection neuronal cells possessed triangular, pyramidal Statistical analysis

or cone-shaped somata, while the stellate neuronal cells
possessed round, oval, spherical or circular shaped
somata (Fig. 6). The hyperpallium apicale region contains
homogenously distributed, purple blue coloured, densely
packed, neuronal somata (Fig. 6A-B). The hippocampus
(Fig. 6C) and the area parahippocampalis (Fig. 6D) of
the hippocampal complex were observed to be populated
by the homogenously distributed, purple blue coloured,
densely packed somata of different neuronal cells. In the
dorsolateral forebrain of chick, the intermediate corticoid
subfield contains more or less homogenously distributed,
less-moderate densely packed, purple-blue colored somata
(Fig. 6E). The dorsolateral corticoid subfield contains
uniformly distributed, moderately packed, purple-blue
colored neuronal somata arranged in the narrow strip like
structure of the corticoid complex (Fig. 6F). In present
study, different neuronal cells show variations in their
soma diameter in all the five sub-regions of telencephalon
in both 15 and 30-day-old chick (Table 1).
Statistical analysis of all the data measured for the
soma diameter was done by using parametric, unpaired,
two tailed, t-test with welch’s correction. The results
obtained from statistical analysis revealed that the age-
factor induces remarkable changes in neuronal soma
diameter of 30-day-old chick in comparison to 15-day-
old chick (Table 1). The soma diameter of different
neuronal cells of 30-day-old chicks were compared with
the soma diameter of different neuronal cells present in
15-day-old chicks in posterior-lateral forebrain, dorso-
medial forebrain and dorso-lateral forebrain for significant
or non-significant decrease or increase (Fig. 7-9).
Posterio-lateral forebrain
The outcomes of statistical analysis for the soma
diameter of multipolar neurons of 30-day-old chick
showed a significant (***P<0.001) decrease in HA, IHA,
and HI layers of hyperpallium apicale whereas a non-
significant decrease was observed in HD layer. A
significant decrease was observed in the soma diameter
2120 Adarsh Kumar et al

Fig. 5 : Drawing of stacked serial cresyl-violet-stained dorsal forebrain sections from rostral to caudal pole (A-M) of 15-day-old chick,
Gallus gallus domesticus showing the position of the (APH) area parahippocampalis, the (VW) visual wulst, the (Hp) hippocampus,
the (CI) intermediate corticoid subfield & the (CDL) dorsolateral corticoid subfield. At the caudal most level, the (HCC) hippocampal
complex is only represented by the parahippocampal area (APH) only & the corticoid complex by the dorsolateral corticoid subfield
only. Below each section the distance from the previous section is given from rostral to caudal end. The section (A) is 4440 µm
posterior from the first rostral section of the cerebral hemisphere. Scale bar = 2mm

of the pyramidal neurons of 30-day-old chick present in
all the hyperpallium apicale layers viz. HA (**P<0.01),
IHA (***P<0.001), HI (**P<0.01) and HD
(***P<0.001). The stellate neurons of 30-day-old chick
showed a non-significant decrease in their soma diameter
of HA layer and a significant (***P<0.001) decrease was
observed in their soma diameter in IHA and HI layers;
whereas a non-significant increase in the soma diameter
was observed in HD layer of hyperpallium apicale (Fig.
7A-C; Table 1).
Dorso-medial forebrain
The hippocampal multipolar neurons of 30-day-old
chick (Fig. 8A) showed a significant decrease in their
soma diameter in (dHp) dorsal hippocampus (*P<0.05),
(mHp) medial hippocampus (***P<0.001) and (vHp)
ventral hippocampus (***P<0.001). The pyramidal
neurons of 30-day-old chick showed a significant
decrease in their soma diameter in dHp (**P<0.01) and
mHp (***P<0.001), whereas in vHp a non-significant
increase was observed in their soma diameter (Fig. 8B).
In dHp a significant (***P<0.001) decrease was observed
in soma diameter of stellate neurons of 30-day-old chick
whereas in mHp and vHp a non-significant decrease was
observed in their soma diameter (Fig. 8C). In the area
parahippocampalis of 30-day-old chick (Fig. 8D) a non-
significant increase was observed in the soma diameter
of multipolar neuronal cells, whereas in the pyramidal
and stellate neuronal cells a non-significant decrease in
their soma diameter was observed in comparison to 15-
day-old chick (Table 1).
Dorso-lateral forebrain
A non-significant increase was observed in the soma
diameter of multipolar neurons present in the layer-I of
intermediate corticoid area of 30-day-old chick in
comparison to 15-day-old chick, whereas in layer-II and
layer-III, the multipolar neurons showed a significant
(**P<0.01) and a non-significant decrease in their soma
diameter, respectively (Fig. 9A). The pyramidal neurons
of layer-I showed a non-significant increase in their soma
diameter, and a significant (***P<0.001) decrease in their
soma diameter in both layer-II, and layer-III (Fig. 9B).
The stellate neurons showed a significant increase in their
soma diameter in layer-I (***P<0.001) and layer-II
(*P<0.05), whereas in layer-III a non-significant increase
was observed in their soma diameter (Fig. 9C). In the
dorsolateral corticoid area of 30-day-old chick (Fig. 9D),
 Age affects the neuronal soma diameter in cerebral cortex of 15 and 30-day-old chick 2121

Fig. 6 : Microphotographs (A-F) showing the cresyl violet stained sections of chick, Gallus gallus domesticus forebrain from rostral to
caudal pole. (A) & (B) showing the visual wulst, (C) & (D) showing the hippocampus proper & the parahippocampal area of the
hippocampal complex, (E) & (F) showing the intermediate & the dorsolateral corticoid areas of the corticoid complex. Arrows 1, 2
& 3 showing small, medium & large sized soma in their respective fields. Section (A) is 1480 µm posterior from the first rostral
section of the cerebral hemisphere, while the section (B) is 2720 µm posterior from section (A), section (C) & (D) are 2600 µm
posterior from section (B), while section (E) & (F) are 800 µm posterior from the section (C) & (D). Magnification 400x. Scale bar
= 50 µm.

a significant (***P<0.001) decrease was observed in the
soma diameter of the multipolar, the pyramidal and stellate
neuronal cells in comparison to 15-day-old chick (Table
1).
DISCUSSION
The results of present study evaluate the effect of
age-factor on the soma diameter of diverse telencephalic
neuronal cells in two different age group (fifteen and
thirty-day-old) of male chicks, Gallus gallus domesticus
along with morphological, anatomical, and histological
aspects of telencephalon for clear understanding by using
Cresyl-Violet-staining method. The present study
demonstrated that the smooth, large sized, pear shaped,
chick brain consists of three major subdivisions: the
rhombencephalon (consists of metencephalon,
myelencephalon and pons varolii), the mesencephalon
(consists of optic bigemina and crura cerebri) and the
prosencephalon (consists of rhinencephalon,
telencephalon or cerebral hemisphere and diencephalon).
Telencephalon of forebrain is one pair of large sized, pear
or obtuse triangle shaped pyriform lobes that are
collectively referred to as cerebrum which is composed
of two symmetrically right and left cerebral hemispheres.
These observations were in accordance with reports in
other birds such as Quail (Al-bakri, 2016), Vana raja birds
(Kumar et al, 2016), Vencobb broiler chicken (Gupta et
al, 2016), Pigeon (Al-Nakeeb and Jasim, 2018), Broiler
2122 Adarsh Kumar et al

Fig. 7 : Showing the results of parametric unpaired two tailed t-test for soma diameter of different neuronal classes observed in the visual
wulst of non-stressed (NS) 15- and 30-day-old chick, Gallus gallus domesticus. Hyperpallium apicale (HA), interstitial nucleus of
the hyperpallium apicale (IHA), hyperpallium intercalatum (HI), and hyperpallium densocellulare (HD). Data is significantly
different at levels: **P < 0.01, and ***P < 0.001.

Fig. 8 : Showing the results of parametric unpaired two tailed t-test for soma diameter of different neuronal classes observed in the
hippocampus (A, B, C) and parahippocampus (D) of non-stressed (NS) 15-, and 30-day-old chick, Gallus gallus domesticus. dorsal
hippocampus (dHp), medial hippocampus (mHp) and ventral hippocampus (vHp). Data is significantly different at levels: *P <
0.05, **P < 0.01, and ***P < 0.001.
 Age affects the neuronal soma diameter in cerebral cortex of 15 and 30-day-old chick 2123

Fig. 9 : Showing the results of parametric unpaired two tailed t-test for soma diameter of different neuronal classes observed in the
intermediate (A, B, C) and dorsolateral corticoid area (D) of non-stressed (NS) 15-, and 30-day-old chick, Gallus gallus domesticus.
Data is significantly different at levels: *P < 0.05, **P < 0.01, and ***P < 0.001.

chicken (Kumaravel et al, 2019) and Ostrich (Koushafar
and Mohammadpour, 2019). The present study also
showed that all the telencephalic external cortical areas
were populated with three types of neuronal cells: the
multipolar, the stellate and the pyramidal neurons on the
basis of soma shape that show homology with the results
of above-mentioned avian studies only in respect of
hippocampal morphology.
The histo-morphological avian studies have revealed
that the wulst is a laminated bulge that occupies the
rostrodorsal surface of the cerebral hemisphere.
Functionally, the wulst has been divided into two subfields:
a small rostral somatosensory motor region and a large
caudal posterolateral visual region known as visual wulst
(VW). The visual wulst is further consists of four
morphologically distinguished, dorso-ventrally arranged
cellular laminae: the hyperpallium apicale (HA), the
interstitial nucleus of the hyperpallium apicale (IHA), the
hyperpallium intercalatum (HI), and the hyperpallium
densocellulare (HD). Towards caudal end, the IHA, the
HI and the HD layers of the visual wulst vanish thus
only the HA layer leftovers, limited ventrally by the
mesopallium in Strawberry Finch (Chand et al, 2013),
pigeon (Shimizu and Karten, 1990), Chick (Deng and
Rogers, 2000, 2002), Zebra finch (Watanabe et al, 2011),
Baya weaver (Srivastava and Gaur, 2013), which is also
observed in present research study.
Different studies were in agreement with the findings
of present study that disclosed the dorsomedial forebrain
is relatively a narrow, curved band of tissue that lies on
the dorsomedial surface of the cerebral hemisphere and
is separated from the rest part of the hemisphere by a
lateral ventricle (VL). The hippocampal complex is
further comprising two major divisions: (a) ventrally
situated V-shaped structure appeared in the ventral two-
third part of the hippocampal complex ranges from medial
to lateral ventricle is called hippocampus and a dorso-
medial curvature of the telencephalon above the angle of
the lateral ventricle extending towards the rostro-caudal
direction is known as the area parahippocampalis. The
hippocampus proper could be subdivided into medial
(HCm), lateral (HCl) and ventral (HCv) hippocampus.
At the most rostral level, the hippocampal complex is
occupied by a small area that is area parahippocampalis
(APH), limited dorsolaterally by the most superficial layer
of the visual wulst (VW) that is hyperpallium apicale
(HA). The area parahippocampalis seems to be a
transitional region between the hippocampal area and the
adjacent telencephalic areas of the cerebral cortex as
observed in Chicken and Homing pigeon (Tömböl et al,
2000), Pigeon (Vargas et al, 2004), P. Krameri (Srivastava
and Sin, 2012), House crow (Singh et al, 2014 and
Srivastava et al, 2016), Vencobb broiler chicken (Gupta
et al, 2016), Strawberry finch (Srivastava et al, 2007),
Parrot and Koel (Singh et al, 2015), Indian roller (Ojha
and Singh, 2021a, 2021b), Vana raja birds (Kumar et al,
2016) and broiler chicken (Kumaravel et al, 2019).
The present study revealed that the corticoid complex
 2124
Table 1 : Showing both the significant and non-significant results of the Unpaired two tailed t-test (with Welch’s correction) for soma diameter of different neuronal classes observed in the
telencephalic areas of 15 & 30-day-old chick, Gallus gallus domesticus.
Soma Diameter (µm)(Mean ± SEM) Unpaired t-test (with Welch’s correction) Results
Region Neurons
15-day-old 30-day-old Df P tcalculated ttable Significant or
chick chick value value value value Non-significant
HA Multipolar 32.46 ± 0.82 22.96 ± 0.68 17 P<0.0001 8.872 2.11 Significant***
Pyramidal 22.63 ± 0.79 19.15 ± 0.49 15 0.0019 3.760 2.131 Significant**
Stellate 11.07 ± 0.28 10.66 ± 0.03 17 0.3247 1.014 2.11 Non-significant
IHA Multipolar 30.89 ± 0.90 19.91 ± 0.69 16 P<0.0001 9.665 2.12 Significant***
Pyramidal 25.97 ± 0.78 15.35 ± 0.42 13 P<0.0001 12.02 2.16 Significant***
Stellate 11.73 ± 0.42 8.05 ± 0.31 16 P<0.0001 7.100 2.12 Significant***
HI Multipolar 28.79 ± 0.72 21.92 ± 0.16 14 0.0002 5.039 2.145 Significant***
Pyramidal 22.21 ± 0.78 19.30 ± 0.47 14 0.0068 3.174 2.145 Significant**
Stellate 9.76 ± 0.30 7.31 ± 0.34 17 P<0.0001 5.379 2.11 Significant***
HD Multipolar 22.56 ± 0.91 21.10 ± 0.66 16 0.2108 1.304 2.12 Non-significant

Adarsh Kumar et al
Pyramidal 22.25 ± 0.94 16.90 ± 0.29 10 0.0003 5.452 2.228 Significant***
Stellate 9.64 ± 0.37 10.34 ± 0.28 16 0.1521 1.504 2.12 Non-significant
dHp Multipolar 22.46 ± 0.89 19.72 ± 0.73 17 0.0292 2.381 2.11 Significant*
Pyramidal 18.72 ± 0.62 16.14 ± 0.21 10 0.0028 3.925 2.228 Significant**
Stellate 12.45 ± 0.47 9.26 ± 0.40 16 P<0.0001 5.160 2.12 Significant***
mHp Multipolar 24.53 ± 0.76 18.04 ± 0.51 15 P<0.0001 7.106 2.131 Significant***
Pyramidal 19.97 ± 0.50 14.50 ± 0.74 15 P<0.0001 6.116 2.131 Significant***
Stellate 9.14 ± 0.22 8.64 ± 0.19 17 0.1034 1.721 2.11 Non-significant
vHp Multipolar 23.86 ± 1.03 18.22 ± 0.49 12 0.0003 4.950 2.179 Significant***
Pyramidal 14.50 ± 0.74 18.95 ± 0.38 16 0.3054 1.059 2.12 Non-significant
Stellate 9.88 ± 0.41 8.77 ± 0.35 17 0.0559 2.052 2.11 Non-significant
APH Multipolar 22.69 ± 0.85 23.30 ± 0.53 15 0.5526 0.6075 2.131 Non-significant
Pyramidal 17.61 ± 0.72 17.25 ± 0.59 17 0.7056 0.3842 2.11 Non-significant
Stellate 9.36 ± 0.58 8.64 ± 0.41 16 0.3288 1.007 2.12 Non-significant
L-I Multipolar 18.43 ± 2.03 26.63 ± 0.26 10 0.2407 1.247 2.228 Non-significant
Pyramidal 16.16 ± 0.63 18.11 ± 0.72 17 0.0571 2.041 2.11 Non-significant
Stellate 7.83 ± 0.15 10.50 ± 0.29 13 P<0.0001 8.278 2.16 Significant***

Table 1 continued...
 Age affects the neuronal soma diameter in cerebral cortex of 15 and 30-day-old chick 2125

Hyperpallium apicale (HA), interstitial nucleus of the hyperpallium apicale (IHA), hyperpallium intercalatum (HI), hyperpallium densocellulare (HD), medial hippocampus (HCm), lateral
hippocampus (HCl), ventral hippocampus (HCv), area parahippocampalis (APH), layer-I (L-I), layer-II (L-II), layer-III (L-III), and dorsolateral corticoid area (CDL). Data is significantly
is present towards the dorsolateral side of the cerebral
cortex and includes two subfields: the dorsolateral
corticoid subfield and the intermediate corticoid subfield.

Non-significant

Non-significant
Significant***

Significant***

Significant***
Significant***
Significant***
Significant** The intermediate corticoid (CI) region is found to be

Significant*
further composed of three layers: Layer-I (present
towards the dorsal surface), Layer-II (present in the
centre of the intermediate corticoid region) and Layer-
III (present beneath the layer-II, towards the lateral
ventricle). At the caudal end of cerebral hemisphere, the
intermediate corticoid area along with the different
subfields of the hippocampal formation progressively
2.131
2.145

2.179
2.179 extinct. Thus, caudally at the level of cerebellum, only
2.16
2.11
2.16
2.12

2.12
the parahippocampal area of the hippocampal complex
and the dorsolateral corticoid area of the dorsolateral
forebrain are present. The dorsolateral corticoid subfield
is devoid of layering pattern and was observed to be a
narrower thin strip-like organization. These findings, of
3.669
4.195
2.767
2.056
5.600
1.531
5.374
4.765
11.48

the present study are in line with those of other studies

conducted in different birds like Parrot (Singh et al, 2019),
Chick and Pigeon (Tömböl et al, 2000b), Zebra finch
(Montagnese et al, 1996), chick, Gallus domesticus
(Kumar et al, 2021 and Kumar et al, 2021), Indian house
P<0.0001

P<0.0001

crow (Srivastava et al, 2014) and strawberry finch

0.0023
0.0009
0.0160
0.0554

0.1452
0.0002
0.0005

(Srivastava et al, 2009).

Additionally, the specific results obtained in the
present study related to the variations induced by age-
factor in soma diameter of diverse neuronal cells observed
in the telencephalon of Gallus gallus domesticus cannot
be related with that of any other bird, because no exact
similar findings on variations induced by age-factor were
15
14
13
17
13
16
12
12
16

available to compare with the results of the present study.

The decrease in neuronal soma diameter of 30-day-old
chick in comparison to 15-day-old chick as shown in the
19.68 ± 0.50
16.45 ± 0.52
10.06 ± 0.18
19.41 ± 0.85
16.16 ± 0.76

19.32 ± 0.91
17.77 ± 0.42
9.79 ± 0.25

7.36 ± 0.32

present study may be because the neurons extend their

dendrites to form new connections in response to external
environment. Thus, the work of soma cytoplasm is
different at levels: *P<0.05, **P<0.01 and ***P<0.001.

extended towards the dendrites and so the soma diameter

constricts with increasing age in presently studied bird,
Gallus gallus domesticus.
22.98 ± 0.75
20.60 ± 0.84

22.07 ± 0.98
20.97 ± 0.39

30.60 ± 0.90
22.40 ± 0.88
13.56 ± 0.44
8.94 ± 0.37

9.14 ± 0.35

CONCLUSION
The findings of the present research study provide a
comparative description of variations in neuronal soma
diameter of various neuronal classes: the multipolar, the
pyramidal, and the stellate neurons in all the external
cortical areas of telencephalon of 15 and 30-day-old
Multipolar

Multipolar

Multipolar
Pyramidal

Pyramidal

Pyramidal

chicks. The data shows that neuronal soma diameter, in

Table 1 continued...

Stellate

Stellate

Stellate

all the regions of 30 day-old-chick, is decreasing whether

it is a significant or non-significant with few exceptions.
Thus, it can be predicted that the soma diameter is
decreasing in response to age factor in all the external
CDL
L-III
L-II

cortical areas of telencephalon. The present research

2126 Adarsh Kumar et al
study might serve as a base for further exciting research
required for finding the relationship of different neurons
with behavior of birds by using some more advanced
techniques for tissue section preparation and staining of
the brain tissue.
Competing interests
The authors declare that they have no competing
interests.
Consent for publication
Not applicable.
Ethics approval
The present research work has been approved by
the Institutional Animal Ethics Committee (IAEC) with
Protocol Number KUDOPS / 106 and all the experimental
procedures performed in the present research study were
carried out strictly in accordance with the animal care
guidelines of the Animal Ethics Committee of Kumaun
University, Nainital. Uttarakhand, India.
Funding
The authors have no financial or personal relationship
with any third party whose interests could be positively
or negatively influenced by the article’s content. This
research did not receive any specific grant from funding
agencies in the public, commercial, or not-for-profit
sectors.
Availability of data and materials
We have shared new data in this research article.
The data that support the findings of this study will be
available from the corresponding author upon reasonable
request.
Author’s contributions
Dr. Ram Chandra Maurya: Conceptualization, Data
Curation, Formal Analysis, Methodology, Project
Administration, Resources, Software, Supervision,
Validation, Writing-Review & Editing. Adarsh Kumar:
Investigation, Methodology, Visualization, Writing-Original
Draft preparation, Writing-Review & Editing. Kavita
Tamta: Writing-Review & Editing; Hemlata Arya:
Writing-Review & editing.
ACKNOWLEDGEMENTS
The authors thank the Head, Department of Zoology
(DST-FIST Sponsored), Soban Singh Jeena Campus
Almora, Kumaun University Nainital for providing
essential infrastructural support for the present research
work.
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Abbreviations
APH, Area parahippocampalis; CC, Corticoid complex; CDL,
Dorsolateral corticoid area; CI, Intermediate corticoid area; dHP,
Dorsal hippocampus; DPX, Dibutylphthalate polystyrene xylene;
HA, Hyperpallium apicale; HD, Hyperpallium densocellulare; HF,
Hippocampal formation; HI, Hyperpallium intercalatum; HP,
Hippocampus; IHA, Interstitial nucleus of the hyperpallium apicale;
mHP, Medial hippocampus; RT, Room temperature; vHP, Ventral
hippocampus; VL, Lateral ventricle; VW, Visual wulst