Dimensions of Cerebral Volume, Prefrontal Cortex, Thalamus, Hippocampus and Varolio Pons, in Bullfighting Bulls and Beef Cattle - Dimensiones del Volumen Cerebral, Corteza Pre-frontal, Tálamo, Hipocampo y Puente de Varólio en Toros de Lidia y en Bovinos Productores de Carne

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org/revistas/redvet
Revista electrónica de Veterinaria - ISSN 1695-7504
Dimensions of Cerebral Volume, Prefrontal Cortex, Thalamus,

Hippocampus and Varolio Pons, in Bullfighting Bulls and Beef Cattle -
Dimensiones del Volumen Cerebral, Corteza Pre-frontal, Tálamo,
Hipocampo y Puente de Varólio en Toros de Lidia y en Bovinos Productores
de Carne
Gouveia, Augusto J.1* Martins, Vítor C.2 Alexandre-Pires G.3
1
Instituto Nacional Investigação Agrária e Veterinária, Av. da República,
2780-157 Oeiras. Portugal. *Corresponding author: Email -
augusto.gouveia@iniav.pt
2
Unidade Estratégica de Investigação de Sistemas Agrários, Florestais e
Sanidade Vegetal. Av. da República, INIAV, 2780-157 Oeiras, Portugal.
3
Departamento de Morfologia e Função; CIISA – Centro de Investigação
Interdisciplinar em Sanidade Animal; Faculdade de Medicina Veterinária,
Univ. de Lisboa. Av. da Universidade Técnica,1300-477 Lisboa, Portugal.
Summary
The Bullfighting Bull (BB) is aggressive, and its function and phenotype, extraordinarily
different from the Beef Cattle Breeds (BCB). It is considered that BB have a closer
proximity to the Bos primigenius [37].
It is accepted that the first taurine activity in Portugal dates from 1258 [18].
The BCB’s suffered meanwhile, genetic and nutritional changes that converged to their
specificity of high productivity and docility.
The divergence is so pronounced that it is important to know if some structures of the
brainstem and limbic system of the BB are parallel with correspondent structures BCB,
since phylogenetically they are associated with the effects of the Archipallium and the
Paleopallium.
Thus, the objective of this study was to compare the dimensions of the Brain Volume,
height of the Prefrontal Cortex and the areas of the Thalamus, Hippocampus and Varolio
Pons in the BB and in BCB.
By the observation of the Components, it is verified that we are in presence of 2 totally
discrepant animals – BB  BCB.
Keywords: Brain Volume | Prefrontal Cortex | Thalamus | Hippocampus | Varolio Pons |

Brain | Comparative Neuroanatomy.
Resumen
Los toros de la Raza Brava de Lidia (BB) son, por su función y fenotipo,
extraordinariamente diferentes de los Bovinos Productores de Carne (BCB); el toro de
lidia es considerado uno de los más cercanos al Bos primigenius [37].
Se admite que la primera actividad taurina en Portugal data de 1258 [18].
Dimensions of Cerebral Volume, Prefrontal Cortex, Thalamus, Hippocampus and Varolio Pons, in Bullfighting Bulls 1
and Beef Cattle
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Los BCB, sin embargo, han sufrido cambios genéticos y nutricionales que han
convergido a su especificidad de alta productividad y docilidad.
La divergencia es tan pronunciada que es importante saber si ciertas estructuras del
tronco encefálico y el sistema límbico de los BB son paralelas a los BCB, ya que están
asociadas filogenéticamente con Archipallium y Paleopallium.
Por lo tanto, el objetivo fue comparar las dimensiones del volumen cerebral, la altura de
la Corteza Pre-frontal y las áreas de Tálamo, Hipocampo y Puente de Varólio en BB y
BCB.
A partir de la observación de los Componentes, se verifica que estamos en presencia de
2 animales totalmente discrepantes – BB  BCB.
Palabras Claves: Volumen Cerebral | Corteza Pre-frontal | Tálamo | Hipocampo | Puente

de Varólio | Cerebro | Neuroanatomía Comparativa.
I Introduction
The brain is one of the most intricate, complicated and impressive evolutionary organ.
The studies of structure / function relationships are fundamental so that we can
understand the evolutionary changes that have occurred [31].
The expansion of the prefrontal cortex is an important evolutionary landmark resulting

naturally in the enlargement of the brain [25]. Phylogenetic changes in the brains occur
only by changes from ancestral ontogeny [24].
The brain and body of animals respond in a similar way as the human brain and body
when threatened [10] Analogies in components of aggression; humans fight in situations
that pre-determine while animals struggle in situations where no one else would fight.[23]
Domestication causes are responsible for changes in brain size in many species, so the
size and volume of the brain were significantly related to the level of domestication. [7] All
behavior is based on the experience and "intelligence" of the individual organism, and
instinctive behavior can be explained as such [39].
Noteworthy invariants have been known in neuroanatomy, such as the proportionality

between neuronal density and the inverse of the cubic root of the brain volume [6].
If brain growth excessively, performance will be lower, limiting any improvement in

cognitive power [19]. Evidence suggests that as the volume of the brain increases, the
number of neurons also increases, but the total density of neurons decreases as a result
of the boost of the density of neural connections [36].
Numerous studies have supported an association between brain abnormalities and

aggressive behavior [15].
There is evidence of an association between general cerebral dysfunction and aggressive

behavior attributable to frontal lobe dysfunction [17]. Aggressive individuals may reflect a
hemispheric asymmetry, with the left prefrontal cortex being typically involved in emotions
[29]. Reductions in the frontal areas of the brain and connected regions are associated
and Beef Cattle
with violence and aggression.[3] It is not surprising that morphometric changes of the
frontal lobe have been proposed to subjugate antisocial behavior in individuals[32]. The
strong activations of the orbitofrontal cortex receive a rich innervation of all major limbic-
hippocampal circuits [28] .There is evidence that the hippocampus and the prefrontal
cortex play an evident and unique role in the control of aggressive behaviors [30].
The thalamus is a gray and differentiated substance, with functional connections at the
cortical, subcortical and cerebellar levels being the absence of interthalamic adhesion
found on average in 16.4% of the cases studied [33]. In the thalamus, stimulation of the
dorsomedial and anterior nuclei, were correlated with emotional reactivity in man and
animals.[22] It has been suggested that the regulation of the thalamus in the aggression is
performed through reciprocal connections with the amygdala [1].
The role of the hippocampus in regulating emotional behavior was first pointed out by
Papez, who drew attention to the role of this structure in the increase in emotional reality
[22]. The reduced volume of the hippocampus negatively influences learning and memory
[8]. A significant positive correlation was found between reduced hippocampal
dimensions and reduced short-term memory retention [26]. .Recent theories of
hippocampal function have broadened its spatial domain in the holding of episodic
moments of memory [12]. There is evidence of small hippocampal volumes and neuronal
integrity, as well as diminished function, in traumatic stress [35].
The brainstem contributes to the evaluation of sensory information and triggers standard
responses in a differentiated spectrum of emotions [38]. Smaller volumes of the brainstem
are associated with highly aggressive individuals [21]. The structures of the brainstem,
notably the Varolio Pons, emerge as crucial nodes that measure aggressive behavior
[20].
It is suggested that cognitive abilities come from changes at all levels of the central
nervous system, including the brainstem [2].
Limbic structures protrude directly and massively over the hypothalamus and the brain
stem, generating autonomic emotional manifestations [11].
Taking in consideration all the reported aspects and to continue previous morphometric
studies on brain structures, this investigation was conducted to compare the Volume of
the Brain (VB), the height of the Prefrontal Cortex (pFC), and the areas of Varolio Pons
(VP), Thalamus (Th) and Hippocampus (Hc), between Bullfighting Bulls and Beef Cattle
Breeds in order to establish possible differences between them.
II Material and Methods
Between July and September 2017, the brains analyzed were randomly collected from
the Mafra Regional Slaughterhouse, according to the processing and cutting chain, in a
total of 28 brains, N = 14 from BB and N = 14 from BCB, being all of them physiologically
adult and with an average weight of 280 kg and 370 kg, respectively.
Each brain was subjected to several evaluations, namely the VB and the measures of the
aforementioned structures, as evidenced in Figure 1 (Hc, and Th), Figure 2 (VP), pFC
height and Hc in Figure 3.
and Beef Cattle
The VB was determined by volume difference in graduated beakers.
The height of the pFC was measured from the anterior extremity of the knee of the
Corpus Callous to the Frontal Pole of the brain.
VP is located above the bulboprotuberancial groove and below the pontopeduncular

groove, that separates it from the cerebral peduncles [34] whose area was determined by
the formula (B + b) x h / 2, given the trapezoidal configuration.
The areas of Th and Hc respectively located around the 3rd ventricle and below the
corpus callous in the retrocomissural pars, [4] were determined by the formula π.a/2.b/2
[9] given its ellipsoid conformation.
Measurements were made with a digital caliper.
The recorded data was analyzed using SPSS Software (V12) supplemented with the
Microsoft Office Excel spreadsheet with KADDSTAT: Statistical Analysis Plug-In to
Microsoft Excel,[16] where the mean, maximum, minimum and standard deviation were
established for BB and BCB, as well as a comparative “Stem & Leaf” graphic (Boxplot) for
the observed variables and, used the t-Student's test.[27]
Principal component analysis is a multivariate statistical technique that consists of

transforming a set of original variables into another set of variables, called principal
components. These have important properties where each main component is a linear
combination of the original variables, are independent of each other and are estimated for
the purpose of retaining the maximum information in terms of the total variation contained
in the source data. When the first main components, usually the 1st and 2nd component
cumulatively hold a high amount of the total information then this profile determined by
this set of variables can be represented by these two new variables, which makes
possible their representation as points in a two-dimensional graphic. Graphically the
operation can be described as the rotation of points existing in a multidimensional space
giving rise to axes, or main components that arranged in a space of two dimensions
represent sufficient variability that could indicate a pattern to be interpreted [5]. In this
representation, the nearer or farther the points are, the greater will be their similarities or
dissimilarities. Taking this in consideration, the method can be used as an instrument for
the classification and grouping of variables and individuals.
Power analysis and data processing: based on an experimental design with 2 groups (BB
and BCB), considering an experimental variability of 15% (β value of 0.15 producing a
central power of 0.85) and considering as modifications more than 20% of the mean
values of each group with a confidence level of 95% (a value of 0.05), the need for a
minimum of 9 animals per group is estimated.
In any case, 14 BB animals and 14 BCB animals (+ 55.5%) were considered.
and Beef Cattle
Figure 1
BCB
A – Hippocampus; B – Thalamus.
Figure 2
BCB
Varolio Pons
and Beef Cattle
Figure 3
BB
A – Prefrontal Cortex Height. B – Hippocampus. CC- Corpus Callosum
The results obtained in the measurements made to VB, pFC, VP, Th and Hc, considering
the average, maximum and minimum as well as the standard deviation for BBs and BCBs
are those presented in Tables 1 and 2.
Table 1
Statistics of 2 Groups
Structures N Average S. D.
BB 14 390,00 31,256
VB
BCB 14 449,64 36,293
BB 14 10,70 0,8691
pFC
BCB 14 22,33 5,4980
BB 14 297,3 29,829
VP
BCB 14 436,1 85,960
BB 14 144,4* 32,760
Th
BCB 14 139,7* 19,199
BB 14 37,00 10,256
Hc
BCB 14 17,60 03,711
* There are no significant differences between both
Table 2
Amplitudes of the considered structures
Structures Maximum Minimum
BB 420 330
VB
BCB 540 400
BB 12,2 9,4
pFC
BCB 31,0 14,3
BB 361,3 259,8
VP
BCB 631,9 251,0
BB 206,7 106,8
Th
BCB 168,1 112,3
Hc BB 62,9 23,4
and Beef Cattle
BCB 24,4 12,3
Boxplot and T-Test for the 5 variables per se (VB, pFC, VP, Th and Hc).
Volume of the Brain - VB:
m3
VB
BB CB
Figure 4
Comparative diagram of the relationship between brain volumes in BBs and BCBs
Highly Significant Difference – 0,001 (0.1%).
Average = 390 cm3 for BB and 450 cm3 for BCB.
In the case of the brain, there is a lower volume for BBs than BCBs. The 95% confidence
interval ranges from 372 to 408 for BB and 429 and 471 for BCBs.
The "Stem & Leaf" graphic (Boxplot) for brain volume shows the difference between
averages, as well as the existence of 4 disparate values such as the cases of the BB No
2, 9 and 12 and, the BCB No 26.
Prefrontal Cortex Height - pFC:
and Beef Cattle
mm
pF
C
BB BCB
Figure 5
Comparative diagram of the relationship between height of the Prefrontal Cortex in BBs and BCBs
Highly Significant Difference – 0,001 (0.1%).
Average = 10.7 mm for BBs and 22.3 mm for BCBs.
In the case of the prefrontal cortex, there is a smaller height for BBs than BCBs. The 95%
confidence interval varies between 10.2 and 11.2 for BBs and 19.2 and 25.5 for BCBs.
The "Stem & Leaf" graphic (Boxplot) for the height of the prefrontal cortex shows the
difference between averages as well as the greater dispersion of values in the case of
BCBs (minimum of 14.3 and maximum of 31.0).
Area of the Varolio Pons – VP:
and Beef Cattle
mm2
VP
RB
BL BPC
BPC
BCB
Figure 6
Comparative diagram of the relationship between the areas of the Varolio Pons in the
BBs and in the BCBs
Highly Significant Difference – 0.001 (0.1%).
Average = 297.3 mm2 for BB and 436.1 mm2 for BCB.
In the case of the Varolio Pons there is a smaller area for BB in relation to the BCBs. The
95% confidence interval varies between 280 and 315 for BBs and 387 and 486 for BCBs.
The "Stem & Leaf" graphic (Boxplot) for the Varolio Pons area shows the difference
between averages as well as the greater dispersion of values in the case of BCBs
(minimum of 251 and maximum of 632).
Thalamus Area – Th:
and Beef Cattle
mm2
Th
BBBLB
R B C
BPCCB
B
Figure 7
Comparative diagram of the relationship between thalamus areas in BBs and BCBs
No Significant Difference – 0.0058 (0.58%).
Average = 144.4 mm2 for BBs and 139.7 mm2 for BCBs
In the case of the thalamus, there is similarity between BBs and BCBs.
The 95% confidence interval varies between 126 and 163 for BBs and 129 and 151 for
BCBs.
The "Stem & Leaf" graphic (Boxplot) for the Th area shows fairness between the
averages as well as the greater dispersion of values in the case of BCB (minimum of 112
and maximum of 168) and the appearance of spurious values related to BB 1, 4, 5 and 8).
Hippocampus Area – Hc:
and Beef Cattle
mm2
Hc
RBBB
L BBCB
PC
Figure 8
Comparative diagram of the relationship between the hippocampus areas in BBs and
BCBs
Highly significant difference – 0.001 (0.1%).
Average = 36.9 mm2 for BB and 17.6 mm2 for BCB.
In the case of the hippocampus, there is a larger area for BBs than the BCBs.
The 95% confidence interval varies between 31.7 and 43.6 for BB and 15.5 and 19.7 for
BCB.
The "Stem & Leaf" graphic (Boxplot) for the hippocampus area shows the difference
between averages as well as the greater dispersion of values in the case of BB (minimum
of 23.4 and maximum of 62.9).
Principal Components Analysis
For the analysis, the variables of Brain Volume (VB), Prefrontal Cortex height (pFC),
Varolio Pons area (VP), Thalamus area (Th) and Hippocampus area (HC) all considered.
were normalized (average - 0 and standard deviation - 1).
and Beef Cattle
Table 3
KMO and Bartlett’s Test
Kaiser-Meyer-Olkin Measure of Sampling Adequacy 0,746
Bartlett’s Test of Sphericity
Approx. Chi-Square 52,914
df 10
Sig. 0,000
In this Table we present the values related to the Kaiser-Meyer-Olkin and Bartlett tests for
the sphericity of the data.
The Kaiser-Meyer-Olkin test value of 0.75 shows the sufficient adequacy of principal
component analysis in the data study and the Bartlett 's test for sphericity where it is
hypothesized that its correlation matrix is a identity matrix is highly significant - 0.001
(0.1%).
Small values of the level of significance indicate that the procedure is useful for data
analysis.
In the analysis of main components, the components (vectors) are linked to the
information under analysis (set of variables considered trying to condense this information
as much as possible.
Table 4 shows that the variance explained by the 1st and 2nd component is 79.5% and
the 1st component represents 56.7% of the variance while the 2nd component represents
22.9%.
Table 4
Total Variance Explained
Table 5 represents scores ranging from -1 to 1 and evaluates the link between the main
component and the variables.
The first main component is linked to the variables pFC height, VP area, Hc area (in the
negative form) and VB (with a slightly lower value). The second component binds to the
Th variable.
and Beef Cattle
Table 5
Rotated Component Matrix
Component
1 2
VB 0,730 0,475
pFC 0,908 0,054
VP 0,864 -0,123
Th -0,066 0,957
Hc -0,841 0,054
Figure 9 shows the projections of the variables Brain Volume (VB), Prefrontal Cortex
height (pFC), Varolio Pons area (VP), Thalamus area (Th) and Hippocampus area (Hc)
in the 1st and 2nd component.
Th
VB
Hc pFC
VP
Figure 9
Component Plot in Rotated Space
The representation relative to the 1st component shows the opposition between the Hc
area and the VB, the height of the pFC and the area of the VP.
Less prominent in the second component, the area of Th opposes the area of Hc, at the
height of the pFC and the area of the VP.
In other words, the first component is connected, gathering information about the
variables VB, pFC, VP and Hc. In this component VB, pFC and VP obtain positive
representation while Hc obtains negative representation.
and Beef Cattle
The second component connects to the variable Th that has the opposition of VP.
Figure 10 shows the projection of the values for the 1st and 2nd components.
RBL
BPC
Figure 10
Comparative Analysis of Components
With this representation, in the 1st component, the BBs get all negative values and so are
represented on the left side of the graph which is in opposition to the BCBs that get
positive values and so are represented in the right graphic side. The observation of this
aspect shows a clear separation between the BBs and the BCBs where, as mentioned
the first two components together represent almost 80% of the information and this
representation of the individuals, as a whole, exposes the significant differences observed
for most variables considered individually in the case of t-Test (note that only the
difference between the areas of the Th is not significant).
The BB No 4 has a value (206.7 mm2) for the area of Th different of all the others and,
this difference is greater for the cases of BBs No 1, 12, H, G and E.
and Beef Cattle
IV Discussion and considerations
Studies on brains in cattle are rare, however, it has been determined that the average
area of the cerebral amygdala in the BB (0.7cm2) is 42% smaller than that of the BCB
(1,2cm2) and that the average weight of the encephalon is respectively of 353 g and 428
g. It was also observed that the frontal poles were narrow in BB and rounded in the BCB
as well as the height of the cerebral cortex at the level of the basal ganglia, which is larger
in the BCB brains when compared with BB. These latter present the dorsal surface less
convex (thinning of the cortex), exhibiting the brain, a trapezoidal cone shape contrasting
with the round shape in the BCB [13].
Another study showed differences concerning GABA, a single significantly different

parameter, with averages values of 0.2 mg / dL for BB, and it can be inferred that its
decrease is the basis of pain disinhibition mechanisms, BCB (0.5 mg / dL) where
analgesia increased per se.[14]
Subsequently, this study compared the data for VB, pFC, VP, Th and Hc between BB and
BCB which showed extremely significant differences in all structures except Th.
Our data corroborates studies that allude to the expansion of the prefrontal region as an
evolutionary landmark and that domestication causes changes in brain size and volume
(BCB). On the other the reduction in the frontal areas of the brain is associated with
aggressiveness and also that the reduced volume of Hc affects negatively the memory
(BB). Also, on the brainstem it is attested that it triggers a differentiated spectrum of
emotions, given the opposable behavior of BB and BCB.
Crossing data from the Portuguese Association of Breeders of Bullfighting Bulls

concerning bravery behaviours of the bulls evaluated under the scope of the present
study, the authors point out that might occur a putative correlation with the mentioned
studied structures and the aggressive/brave behaviour in Bullfighting Bulls, which contrast
to the docility of the Beef Cattle Breeds.
Although not mentioned in this study, we can empirically claim other observations
considering morphological differences between BBs and BCBs, these resulting from the
functions attributable to both, emphasizing the shape of the horns and the development of
the anterior third in BB against the growth of the posterior third in the BCBs.
Considering and analyzing all the data and elements, which are extremely different and
stable, we wonder if BBs will be a subspecies whose individuals could be called Bos
taurus bravus.
V Conclusion
In order to achieve the objective, we set out, we consider that for the studied groups, the
collection of data obtained and measured technically and scientifically was demonstrative,
presenting a highly significant statistical level in comparative measurements, with the
exception of the thalamus.
This work of morphological parameterization of the brain and brain structures, allowed to
conclude that on average Cerebral Volume, the height of the Prefrontal Cortex and the
and Beef Cattle
area of the Varolio Pons are quite smaller in the Bullfighting Bulls and that the
Hippocampus area, also on average, is much lower in the Beef Cattle Breeds. The
thalamus has a similar weighted area.
From the analysis of Components, it is recognized that between Bullfighting Bulls and
Beef Cattle Breeds there is a clear and sharp separation. Considering these facts, in
comparison, we may be facing neuroanatomical divergent animals.
Acknowledgments
 To the President of the Board of INIAV, I.P. Doctor Nuno Canada.
 To Dr. Eurico Esteves, Administrator of the Regional Slaughterhouse of Mafra.
 To Dr. Vasco Lucas and to the Portuguese Association of Bullfighting Bulls
Breeders.
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Dimensions of Cerebral Volume, Prefrontal Cortex, Thalamus, Hippocampus and Varolio Pons, in Bullfighting Bulls and Beef Cattle - Dimensiones del Volumen Cerebral, Corteza Pre-frontal, Tálamo, Hipocampo y Puente de Varólio en Toros de Lidia y en Bovinos Productores de Carne

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Dimensions of Cerebral Volume, Prefrontal Cortex, Thalamus, Hippocampus and Varolio Pons, in Bullfighting Bulls and Beef Cattle - Dimensiones del Volumen Cerebral, Corteza Pre-frontal, Tálamo, Hipocampo y Puente de Varólio en Toros de Lidia y en Bovinos Productores de Carne

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Dimensions of Cerebral Volume, Prefrontal Cortex, Thalamus,

Keywords: Brain Volume | Prefrontal Cortex | Thalamus | Hippocampus | Varolio Pons |

Palabras Claves: Volumen Cerebral | Corteza Pre-frontal | Tálamo | Hipocampo | Puente

The expansion of the prefrontal cortex is an important evolutionary landmark resulting

Noteworthy invariants have been known in neuroanatomy, such as the proportionality

If brain growth excessively, performance will be lower, limiting any improvement in

Numerous studies have supported an association between brain abnormalities and

There is evidence of an association between general cerebral dysfunction and aggressive

II Material and Methods

The VB was determined by volume difference in graduated beakers.

VP is located above the bulboprotuberancial groove and below the pontopeduncular

Measurements were made with a digital caliper.

Principal component analysis is a multivariate statistical technique that consists of

In any case, 14 BB animals and 14 BCB animals (+ 55.5%) were considered.

BCB 24,4 12,3

Volume of the Brain - VB:

Highly Significant Difference – 0,001 (0.1%).

Average = 390 cm3 for BB and 450 cm3 for BCB.

Prefrontal Cortex Height - pFC:

Highly Significant Difference – 0,001 (0.1%).

Average = 10.7 mm for BBs and 22.3 mm for BCBs.

Area of the Varolio Pons – VP:

Highly Significant Difference – 0.001 (0.1%).

Average = 297.3 mm2 for BB and 436.1 mm2 for BCB.

Thalamus Area – Th:

No Significant Difference – 0.0058 (0.58%).

Hippocampus Area – Hc:

Highly significant difference – 0.001 (0.1%).

Average = 36.9 mm2 for BB and 17.6 mm2 for BCB.

Principal Components Analysis

Comparative Analysis of Components

IV Discussion and considerations

Another study showed differences concerning GABA, a single significantly different

Crossing data from the Portuguese Association of Breeders of Bullfighting Bulls

1. Andy, O., J. Giurintano, L. Giurintano, S. and. McDonald, T. (1975). Modulation of

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