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European Journal of Applied Sciences – Vol. 13, No. 1
Publication Date: February 25, 2025
DOI:10.14738/aivp.131.18314.
Tulp, O. L. (2025). Impact of Brown Adipose Tissue Reduction and Overfeeding on Adiposity and Depotspecific Adipose Tissue
Cellularity: Brown Fat Reduction and Adipose Tissue Cellularity. European Journal of Applied Sciences, Vol - 13(1). 397-408.
Services for Science and Education – United Kingdom
Impact of Brown Adipose Tissue Reduction and Overfeeding on
Adiposity and Depotspecific Adipose Tissue Cellularity: Brown
Fat Reduction and Adipose Tissue Cellularity
Orien L Tulp
ORCID: 0000-0001-6904-2573
Colleges of Medicine and Graduate Studies,
University of Science Arts and Technology,
Montserrat, British West Indies, MSR1110
ABSTRACT
Brown adipose tissue contributes to adaptive changes in metabolic energy
expenditure in response to alterations in diet and environmental conditions,
thereby assisting an animal to maintain thermoregulation and energy balance in
various mammalian species including man and animals. Cafeteria overfeeding of
normally lean rats during early postweaning growth typically results in significant
hyperplasia and in an increased capacity non-shivering thermogenesis and energy
expenditure in brown adipose tissue (BAT). The Interscapular BAT depot is readily
surgically accessible and normally represents approximately one third of the total
BAT mass in lean Sprague Dawley (SD) rats. The effects of experimental
overnutrition via offering a Cafeteria feeding regimen (Café) combined with
surgical reduction of the IBAT mass of adipose tissue cellularity and regional fat
deposition was determined in lean SD rats during 8 weeks of postweaning growth
and development to adulthood. Groups (n= 8 rats/group) of male, SD rats were fed
a Purina Chow diet or the Chow diet plus the Café regimen for 52 days from weaning.
An additional group of the Chow+Café regimen were subjected to surgical removal
of their IBAT at 4 weeks of age and continued on the Chow+Café thereafter (Café- IBAT). At 80 days of age, measures of adiposity including anthropometrics and the
mass and adipocyte cellularity in principle abdominal and subcutaneous fat depots
were determined. Body weight (BW) and mid-abdominal girth were ~20% greater
in Café and Café-IBAT, while linear growth was similar in all groups. The mass of all
fat depots was greater in Café fed animals (p=<0.05) and increased further in
subcutaneous depots and total WAT accumulation in the CaféIBAT animals
(p=<0.05). Adipocyte lipid content and cell diameter of Café > controls in all depots
with further increases in abdominal depots with Café-IBAT. Adipocyte number per
WAT depots of Café > Control in all depots, with further increases in the Inguinal SC
depot. Thus, these results are consistent with regional differences in the effects of
Café feeding on postweaning adipose tissue hyperplasia, hypertrophy and depot
mass and which underwent additional depot-specific differentiation in Café-IBAT.
In conclusion, Café resulted in adipocyte hypertrophy in all depots studied, but the
partial reduction of BAT mass in Cafe-IBAT rats resulted in only modest additional
impact on overall adiposity during overfeeding, with the greatest impact in the ING
SC depot, and thereby consistent with potential thermogenic compensation in other
BAT depots to partially minimize the overall impact of the Café overfeeding regimen
on developing adiposity in this strain.
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Keywords: Obesity, Brown Adipose Tissue, Overnutrition, Adipose Cellularity, Adiposity,
Rat.
INTRODUCTION
The magnitude of diet-induced hormonal effects on adipose tissue cellularity and fat accretion
exert differential impacts in subcutaneous vs. abdominal depots.1 The metabolic effects of
brown adipose tissue on the expression of nonshivering thermogenesis (NST) responses to
alterations in diet and environment in rodents are well established, implying a possible role of
BAT on energy balance in mammalian species by dissipating excess caloric intake as heat while
minimizing energy retention in adipose tissues.2,3 Several authors have reported that surgical
reduction of brown adipose tissue resulted in decreases in the capacity for NST in addition to a
greater propensity to develop greater mass and fatness in white adipose tissue depots,
analogous to that which occurs following pharmacologic ablation.4-8 As noted above, the BAT
has also been proposed as a potential buffer to maintain energy balance against excess weight
gain following episodes of excess caloric intake or macronutrient imbalance.9 In addition,
deficits in BAT energy expenditure secondary to multiple physiological factors have been
observed to occur in the obese phenotype of several rodent strains, where they are likely
metabolic contributors to the excess weight gain and early onset adiposity in those animal
species.1013 The current increasing prevalence of obesity and its numerous pathophysiologic
sequelae in Westernized societies represents a serious challenge to the effectiveness,
availability and capacity of treatment resources to manage the disorders linked to the obesity- linked stigmata.14,15 Adipose tissue is now known to contribute a broad range of hormonal
activities, in addition to the roles of insulin, catecholaminergic, thyroidal, and glucocorticoid
regulation.16-20 The nutritional and metabolic factors that contribute to the current epidemic of
obesity and overweight conditions in Western society typically include factors of diet,
environment, and lifestyle, often combined with links to metabolic and genomic
predispositions.18-20 The therapeutic measures to treat the disorders linked to obesity and
overweight conditions typically address one or more of the adaptive or heritable contributors
but often may be only partially effective due to the complex nature of the conditions.
Brown adipose tissue was discovered to occur in humans and other mammalian species several
centuries ago, but its physiologic role in energy metabolism has been discovered only more
recently.2,3,20-22 The morphology and functions of brown adipose tissue differ markedly from
that of white adipose tissue.15,22-25 Both tissue types occupy specific anatomic domains that are
anatomically suited to their physiologic functional role in homeostasis and energy balance.2,3
In white adipose tissue, the primary function is linked to energy storage in the form of
triglycerides in both abdominal and subcutaneous locations. The triglycerides may be formed
via hepatic or dietary origin, and may become deposited in mature adipocytes or preadipocytes
as a single large lipid droplet, surrounded with a thin layer of cytoplasm, a flattened nucleus
located in the cytoplasmic ring, accompanied with the host of cellular organelles common to
somatic cells, and facilitate the processes associated with our basic cellular functions including
cellular metabolism to maintain cellular viability.23 In times of caloric excess, the lipid droplet
may expand to contain up to 1 μg or more of lipid content in response to hormonal and
substrate influences, while in states of caloric deprivation, the lipid droplet may be mobilized
along its outer surfaces to release free fatty acids, thereby contributing to the energy demands
of peripheral tissues.3,23 Adipocytes from white adipocyte tissue may develop from
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of Applied Sciences, Vol - 13(1). 376-396.
URL: http://dx.doi.org/10.14738/aivp.131.18314
preadipocytes via hyperplasia and hypertrophy and once differentiated, remain viable
thereafter. Adipocytes remain responsive to accommodating increases or decreases in energy
intake and thus serve as an energy reserve from the stored triglycerides and fatty acids
throughout adolescence and much of the adult lifespan. Of significant concern, the onset of
overweight and obese conditions is now occurring earlier in the lifespan and remaining longer
than in previous generations. Overweight and obesity now occur more often in children and
adolescents where it can contribute to an earlier onset of the comorbidities commonly
associated with the overweight and obese conditions.14,15
In contrast, the physiological functions and cellular morphology of brown adipose tissue vary
considerably from those of white AT.2,3,22-24 Brown adipocytes are typically smaller, spherical
structures with a round, centrally placed nucleus and abundant specialized mitochondria. The
main physiological function of brown adipose tissue is to contribute to heat generation and
energy expenditure in response to alterations in diet and environment, accomplished via a large
cytoplasmic density of specialized mitochondria.23,25 BAT depots are strategically located in
close proximity to abundant vascular tissues such that the heat generated my circulate to both
central and peripheral tissues. Thus, brown adipose tissue can effectively dissipate the
endogenous heat energy to peripheral tissues to facilitate the regulation of homeostatic body
temperatures.2,3,23-25 Brown adipocytes can mobilize the heat generation process rapidly, via
specialized β-neuroadrenergic membrane-bound receptors and a dedicated
sympathoneurologic presence.3
Because the lipid in brown adipocytes is contained in multiple small locules distributed
throughout the cytoplasm, it provides a greater net metabolizable surface area to lipid content
ratio. Thus, locular surface area is an important consideration that further contributes to the
efficiency of their energy producing functions. The lipid locules are also strategically
distributed throughout the cytoplasmic compartment in proximity to the specialized
mitochondria, thereby creating a larger metabolizable surface area per unit of lipid than occurs
in white adipocytes. The relatively greater metabolizable surface area thereby facilitates a more
rapid mobilization of the contained lipid since lipid mobilization in both tissue types occur
along the outer surface areas of the lipid droplets or locules. Additionally, because the brown
adipocytes contain a centrally located spherical nucleus, surrounded by all essential organelles
required to maintain cellular functions, the ease of morphologic distinction from other
surrounding cells and tissues is readily discerned with or independently of histological staining
or immunoreactive techniques. Brown adipose tissue develops via hyperplasia and limited
hypertrophy prior to adulthood in the rat, while white adipose tissue in most depots may
continue to increase by hyperplasia from preadipocytes and limited hypertrophy throughout
much of the lifespan in rodents.23,24, Once formed, both brown and white adipocytes appear
remain present thereafter and where they may continue to expand in response to caloric status.
Once formed, differentiated adipocytes of either type can remain active as lipid storage depots
virtually indefinitely, to accommodate the energy needs of the organism as needed during both
energy privation and excess. Both tissues normally regulate their lipid stores and metabolic
status via hormonal actions including those inspired by insulin and catecholamines. In contrast,
pharmacologic ablation via inhibition of β-adrenergic functions results in increases in locule
diameter and lipid content in isolated brown adipocytes, consistent with decreased
thermogenic activity.23
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The primary metabolic roles of Insulin, catecholamines, glucocorticoids, and macronutrient
energy intake regulation in addition to numerous other secondary factors via both direct
stimulatory and permissive effects regulation of energy balance in man and animals.15-18 Insulin
functions exhibit both glycemic and lipogenic actions in peripheral tissues via stimulating
glucose uptake, stimulating lipogenesis and lipid uptake of preformed lipids, while impeding
lipid mobilization and ketogenesis from white adipose tissue stores. Thus, insulin is a primary
and highly influential hormone in the peripheral management of energy stores over time.
Catecholamines can bring about the mobilization of glycogen and lipid stores during instances
of duress, including negative energy balance, macronutrient imbalance, and/or food
depravation. While the metabolic effects of insulin actions may persist for hours, the responses
to catecholamine occur rapidly and are initiated via adrenoreceptor actions located on the
plasma membrane. Accordingly, the positive effects of catecholamines on energy balance are
more immediate and give rise to the initiation of the common ‘fight or flight’ response. Plasma
glucose can become significantly elevated within minutes in response to catecholaminergic
stimulation, while in the presence of insulin resistance, plasma glucose and processes of glucose
disposal in peripheral tissues may persist for hours and take longer to recover after the
adrenergic challenge.26
Brown adipocytes have specialized β3-adenoreceptors, in addition to individual
sympathomimetic neural synapses to facilitate highly specialized and virtually instantaneous
responses once activated .3 In white adipose tissue, insulin typically impedes lipoprotein
mobilization, thereby enzymatically limiting the mobilization of free fatty acids and
diglycerides from stored triglycerides.16 Thus, the overall integrated regulation of energy stores
from carbohydrate and lipid sources is a complex and ongoing process, and which may continue
throughout the healthful lifespan of the individual or animal species baring pharmacologic
inactivation.
The presence of brown adipose tissue has been noted in humans for many generations. Among
the earliest observations were those noted during cadaveric dissections some 1500 years ago.21
More recently, histologic evidence in humans has been demonstrated to occur throughout
much of the lifespan.21,22,27 Thus, while the existence of BAT in mammalian species has been
known for many years, the biochemical processes of energy generation in brown adipose tissue
and its presence in adult humans have been established with advances technologic and
diagnostic processes only more recently.2,3,22,23,25 In rodents, the IBAT depot represents
approximately one third of the total BAT mass, and represents the depot that is most
conveniently accessible from a surgical perspective. Thus, the purpose of the present study was
to determine if partial reduction of brown adipose tissue would result in measurable changes
in adiposity and energy storage in a normally lean rat model of following induced hyperphagia
by offering unlimited access to the cafeteria diet approach, typically consisting of multiple
appetizing, energy rich foods common to the Western diet consumed in many developed
countries.5
MATERIAL AND METHODS
Groups of weanling Sprague-Dawley Rats (n = 6-8 rats/group, 40-42 g BW each) were obtained
from Charles River Laboratories, and acclimated to plexiglass shoebox cages in littermate pairs
and fed Purina Chow and house water, as libitum. At 4 weeks of age, two groups were offered a
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of Applied Sciences, Vol - 13(1). 376-396.
URL: http://dx.doi.org/10.14738/aivp.131.18314
highly palatable Cafeteria diet (Café) in addition to the Purina chow regimen. At 4 weeks of age,
one group of the Café diet regimen was subjected to surgical removal of the entire Interscapular
brown adipose tissue depot (Café-IBAT) under pentobarbitalketamine anesthesia as described
elsewhere 5,7-9 and continued on the Café diet regime thereafter. Recovery from the surgery was
uneventful and complete within a few days. Body weights were monitored periodically
throughout the study. After 52 days of the Café regimens animals were sacrificed via cervical
dislocation and measures of biometry including torso length and girth diameter determined
with an anthropometry tape. The epididymal, retroperitoneal, mesenteric, inguinal and dorsal
adipose tissue depots dissected in their entirety, weighed to the nearest mg, and prepared for
measures of adipose tissue cellularity via the osmium fixation methods of Hirsch and Gallian as
performed in our laboratory.24,27,28 Tissue lipid content was determined gravimetrically via the
microchemical method of Dole and Meinertz as conducted in our laboratory.26,30 Measures of
adipocyte diameters were determined with a stage micrometer via light microscopy at 45X
magnification.23 Data were analyzed by ANOVA corrected for multiple comparisons where
indicated and descriptive analysis via standard statistical procedures.31,32 The study was
approved by the Institutional Animal Care and Use Committee.
RESULTS
The measures of biometry after 52 days of the chow or café regimens are depicted in Figure 1
and 2 and show that the Café diet resulted in significantly greater final body weights in both the
Café and the Café-IBAT groups. There were no significant differences in the initial or final body
weight between the Café and Café-IBAT groups however, suggesting that the ablation of IBAT
likely induced only a modest impact on overall parameters of adiposity. Measures of torso
length were similar in all groups, indicative of an adequacy in macro- and micro- nutrient intake
when fed the Café diet. Measures of mid-abdominal girth and girth to torso length ratio were
greater in the Café fed group and were similar in both Café and Café-IBAT groups.
Figure 1: Effects of diet and IBAT removal on initial and final body weight of rats. Data are the
mean ± 1 SEM, N= 8 rats/group.
0
200
400
600
800
1000
INITIAL BW FINAL BW
p = < 0.05
FIGURE 1: BODY WEIGHTS OF RATS
CONTROL CAFÉ CAFÉ-IBAT
p = n.s.
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Figure 2: Effects of diet and IBAT removal on anthropomentric measures. Data are the mean ± 1
SEM, N= 8 rats/group.
Measures of the mass of principle far depots is depicted in Figure 3, and indicated that the mass
of the abdominal (Epididymal (EPI), Retroperitoneal (RP), and Mesenteric (MES), and the
subcutaneous (Dorsal (DOR), and Inguinal (ING)) fat pads were greater in Café fed than control
rats, with further increases in depot mass in the DOR, ING, and total fat pad mass in the Café- IBAT group. The AT cell number of Café > CHOW in subcutaneous (DOR and ING) and abdominal
RP, and unchanged in EPI. Adipocyte diameter and cellular lipid content of Café > CHOW in all
depots. Cell number increased further in DOR, EPI and RP, and the overall differences in cell
diameter corresponded to the measures of cell lipid content. In addition, surgical reduction of
IBAT resulted in further increases in the mass, cell size and cell lipid content particularly in the
ING depot. In the Café diet, IBAT cellularity determinations resulted in greater depot cell
numbers in the dorsal, retroperitoneal and inguinal depots. The café-IBAT group demonstrated
greater cell numbers in the Café-IBAT inguinal depot, while depot cell numbers in the Café-IBAT
retroperitoneal group were also elevated at a level intermediate between the chow and Café
groups.
0
5
10
15
20
25
30
35
TORSO, cm. GIRTH, cm GIRTH/TORSO x10
p = n .s.
FIGURE 2: BIOMETRY OF RATS
CONTROL CAFÉ CAFÉ-IBAT
*
*
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Figure 3: Effects of diet and IBAT removal on adipose tissue mass. Data are the mean ± 1 SEM,
N= 8 rats/group. Abdominal represent the arithmetic sum of the epididymal, mesenteric and
retroperitoneal depots, and subcutaneous represents the sum of the Dorsal and inguinal
depots. The SUM represents the arithmetic sum of the abdominal and subcutaneous depots,
and the Adiposity represents the sum of the fat pad mass divided by the final body weights of
the rats.
Figure 4: Effects of diet and IBAT removal on adipocyte diameter. Data are the mean ± 1 SEM,
N= 8 rats/group. Abdominal cell diameters represent the arithmetic sum of the epididymal,
mesenteric and retroperitoneal depots, and subcutaneous cell diameters represent the sum of
the mean of the dorsal and inguinal adipocytes. The mean cell diameter represents the
arithmetic mean of the abdominal and subcutaneous depots.
0
50
100
150
200
ABDOMINAL WAT, g SUBCUTANEOUS
WAT, g
SUM WAT ADIPOSITY
p = < 0.05 (Control vs Cafe;)
FIGURE 3: EFFECT OF CAFE' AND IBAT ON ADIPOSITY
CONTROL CAFÉ CAFÉ-IBAT
0
20
40
60
80
100
120
CELL DIA, SQ CELL DIA, ABD MEAN CELL DIA
p = trend (Control vs. Cafe')
FIGURE 4: MEAN ADIPOCYTE DIAMETER
CONTROL CAFÉ CAFÉ-IBAT
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Figure 5: Effects of diet and IBAT removal on adipocyte lipid content. Data are the mean ± 1
SEM, N= 8 rats/group. Abdominal cell lipid content represents the mean of the epididymal,
mesenteric and retroperitoneal depots, and subcutaneous cell lipid content represents the
mean of the dorsal and inguinal adipocytes. The mean cell lipid content represents the
arithmetic mean of the abdominal and subcutaneous depots.
Figure 6: Effects of diet and IBAT removal on adipocyte number. Data are the mean ± 1 SEM, N=
8 rats/group. Abdominal cell lipid content represents the mean cell number/depot of the
epididymal, mesenteric and retroperitoneal depots (right panel), and subcutaneous cell
number (Left panel) represents the mean of the dorsal and inguinal adipocytes. The mean cell
number in the far-right panel represents the arithmetic mean of the cell number in the
abdominal and subcutaneous depots.
0
0.2
0.4
0.6
0.8
1
1.2
CELL LIPID, SQ, ug CELL LIPID, ABD, ug MEAN LIPID, ug
p = < 0.05 (Control vs Cafe')
FIGURE 5: MEAN CELL LIPID CONTENT
CONTROL CAFÉ CAFÉ-IBAT
0
20
40
60
80
100
120
SQ CELL NR ABD CELL NR SUM SQ+ABD
p = < 0.05 (Control vs Cafe')
FIGURE 6: EFFECT OF DIET AND IBAT ON ADIPOCYTE CELLULARITY
CONTROL CAFÉ CAFÉ-IBAT
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of Applied Sciences, Vol - 13(1). 376-396.
URL: http://dx.doi.org/10.14738/aivp.131.18314
DISCUSSION
The presence of brown adipose tissue in hibernating and other animals had previously been
reported for many years, but its relationship to energy balance and diet-induced adaptive
thermogenesis in mammalian species and humans remained unknown or controversial for
many years. 2,3 In the 20th Century, Sims et al were among the first in recent history to describe
the phenomena of ‘luxus consumption’ in humans, which they had observed in human
volunteers in the Vermont Study of Obesity.33 In that study, healthy, normal weight and
predominantly sedentary volunteers consumed up to 10,000 excess calories/day without
becoming obese, and most subjects quickly lost the modest extra weight they had gained upon
return to normal energy intakes and without significant changes in daily physical activity.32
Rothwell and Stock later demonstrated the apparent presence of brown fat activity in humans
following adrenergic activation, thereby further linking the thermogenic response to dietary
and environmental factors.22 While the presence of brown adipose tissue in hibernating
animals and in humans had been known, the biochemical mechanisms implicated and the
physiologic contributions have now been elucidated by Himms-Hagen and others.2,3 The
thermogenic mechanism was found to revolve around specialized mitochondria in BAT that are
capable of generating heat from the hydrolysis of high energy phosphate bonds from ATP, a
process sometimes referred to as ‘uncoupled’ oxidative phosphorylation.2,3 The process results
in the generation of ~ 7 kcal/mole of high energy phosphate bonds as heat and typically occurs
without being linked to biosynthetic processes.3The effects of pharmacologic inhibition of BAT
thermogenesis via β-blockade resulted in microscopic enlargement of the lipid locule diameters
indicative of decreased thermogenic activity.23 Attempts to document excess weight gain and
greater adiposity following surgical removal of IBAT have yielded variable results.5-7, 9,10,24
In conclusion, the results of this study indicate that café- induced overfeeding resulted in depot
specific increases in adiposity including increases in body weight, abdominal circumference,
and in adipose tissue cellularity in abdominal and subcutaneous depots. The results obtained
herein are qualitatively similar to those that have been reported by other authors. 6,7,9,10,14,24,35-
37 In the present study, the increases in adipose tissue cellularity occurred by a combination of
hyperplasia and hypertrophy of adipocytes that were expressed differently in different depots.
In the epididymal depot, adipocyte hyperplasia is likely complete by puberty, and increased in
depot mass beyond puberty occur via limited hypertrophy. Epididymal cell diameters and cell
lipid content exhibited only a modest trend toward greater lipid content following the Café diet
in either Café group or Café-IBAT were without additional effect. Likewise, adipocyte number
in the dorsal, retroperitoneal and inguinal depots increased significantly when fed the café diet,
but the cell number in the Café-IBAT group was greater only in the Inguinal depot. The reasons
for the differential effects of overfeeding on differential depot specific alterations in adipose
tissue cellularity are unclear but may be secondary to depot specific differences in insulin
sensitivity, lipogenic actions, and chronological differences in lipid accretion in the different
depots. Effects on linear growth were not observed, suggesting that the Café diet as offered in
combination with their chow regimen likely provided adequate nutrition to accommodate lean
tissue growth and development. In contrast, the differences in adiposity correlated more with
net energy intake, including refined carbohydrates and processed food items of less nutritional
value. While measures of adipocyte number in the IBAT were not measured in the present
study, parallel studies in both rats and mice reported an approximate 3-fold increase in IBAT
mass and cellularity after an analogous 52-day or shorter feeding regimen.7,8,24,29,34-37
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SUMMARY AND CONCLUSIONS
The results of this study showed that feeding a Café diet supplement to normally lean rat during
the postweaning growth period resulted in depot-specific increases in mass and cellularity
characteristics in lean, Sprague-Dawley rats. Partial surgical reduction of BAT mass by
removing the Interscapular depot in its entirety resulted in modest additional depot-specific
increases in adiposity, but the net increases were not proportionate to the proportion of BAT
removed. Whether other BAT depots may have compensated by additional increases in
cellularity or thermogenic functions remain unclear, however in an earlier study, resting and
norepinephrine simulated thermogenic responses were only modestly decreased following a
similar surgical reduction of IBAT.5,6 Thus, overfeeding via the Café regimen in a strain of
normally lean rats is an effective method to bring about modest physiological changes in
adiposity and adipose tissue cellularity in white adipose tissue depots via differential depot- specific effects on adipocyte hyperplasia and hypertrophy.
ACKNOWLEDGEMENTS
The author thanks the University of Science Arts and Technology, Montserrat for the
Institutional resources to complete this study.
Application of AI (Artificial Intelligence) Disclaimer
Author hereby declares that NO generative AI technologies such as Large Language Models
(ChatGPT, COPILOT, etc) and text-to-image generators have been used during the writing or
editing of this manuscript.
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