Feyisa et al.

Laboratory Animal Research

https://doi.org/10.1186/s42826-019-0024-y
(2019) 35:29
Laboratory Animal Research

RESEARCH Open Access

Investigation of the effect of coffee on

body weight, serum glucose, uric acid and
lipid profile levels in male albino Wistar rats
feeding on high-fructose diet
Teka Obsa Feyisa1* , Daniel Seifu Melka2, Menakath Menon2, Wajana Lako Labisso2 and Mezgebu Legesse Habte1

Abstract
Coffee is one of the most commonly consumed beverages in the worldwide and is assumed to have protective
effects against metabolic syndrome. The present study was aimed at investigating the effect of coffee on body
weight, serum glucose, uric acid and lipid profile levels in male albino Wistar rats feeding on high fructose diet. A
post-test experimental study was conducted on a total of 30 (9–10 weeks old) male albino Wistar rats. The rats were
divided into 6 groups: group I (normal control)-fed on standard chow and plain tap water only; group II (fructose
control)-fed on standard chow and 20% of fructose solution; group III–VI (treatment groups)-fed on standard chow,
20% of fructose solution and treated with 71, 142, 213 and 284 mg/kg body weight/day of coffee respectively for
six weeks. At the end, body weight, serum glucose, uric acid and lipid profile levels were investigated. Data was
entered and cleared by epi-data software version 3.1 and analyzed by one way ANOVA followed by Tukey post hoc
multiple comparison tests using SPSS V. 23.00. Statistical significance was considered at p < 0.05. The results showed
that body weight, fasting serum glucose and uric acid levels significantly lowered in rats treated with 213 (p = 0.047;
0.049; 0.026) and 284 (p = 0.035; 0.029; 0.010) mg/kg body weight/day of coffee compared to fructose control
group. Fasting serum triglycide (TG) and low density lipoprotein (LDL-C) levels showed significant reduction in rats
treated with 284 mg/kg body weight/day of coffee as compared to fructose control group (p = 0.031; 0.046) respectively.
In conclusion, treating rats with coffee decreased body weight, fasting serum glucose, uric acid, TC, TG and LDL-C, and
increased HDL-C in a dose dependent manner in rats feeding on high fructose diet, suggesting that coffee consumption
may be helpful in ameliorating metabolic syndrome.
Keywords: Body weight, Coffee, Glucose, High-fructose diet, Lipid profiles, Uric acid

Introduction though the main physiological effects resulting from its

Background consumption are usually ascribed to the presence of caf-
Coffee is one of the most frequently consumed bever- feine, coffee is also extremely enriched with chlorogenic
ages in the worldwide and its beneficial effects on hu- acids (CGA), melanoidins and diterpenes [3, 4].
man health have become a subject matter of several Coffee is prepared by several methods, which signifi-
scientific studies [1]. There are hundreds of variety spe- cantly affect the aroma, flavor and composition of coffee.
cies of coffee, however, commercially two species are Many of the coffee compounds, responsible for its
mostly available: Coffea arabica, about 70% and Coffea unique taste and smell, are formed during the roasting
canephora [2]. Coffee contains a multitude of chemical process [5]. On the contrary, the roasting process may
substances, many of which are biologically active. Even cause degradation of several other compounds, including
antioxidant polyphenols. Depending on the brewing pro-
* Correspondence: takobsi2008@gmail.com cesses, three major types of coffee can be distinguished:
1
Department of Medical Biochemistry, College of Health and Medical (i) boiled unfiltered coffee, (ii) filtered coffee and (iii) de-
Sciences, Haramaya University, Harar, Ethiopia
Full list of author information is available at the end of the article
caffeinated coffee (DC). The property of each type of

© The Author(s). 2019 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver
(http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.
Feyisa et al. Laboratory Animal Research (2019) 35:29 Page 2 of 8

coffee is related to specific granulation of coffee powder, Therefore, the total sample size was 6 groups × 5
water/coffee ratio, temperature and brewing time [6]. rats = 30 rats.
Many in vitro and in vivo studies, together with
epidemiological and human trials have suggested Inclusion and exclusion criteria
beneficial health effects of coffee. Acute or regular Inclusion criteria were age (9-10 weeks old) and normal
coffee consumption may reduce the risk factors of body weight (< 250 g).
mortality, cardiovascular disease, T2DM, obesity, liver Exclusion criteria were suffering from diarrhea, or any
disease, cancer and many of degenerative diseases observable physical abnormality before treatment.
such as Alzheimer’s and Parkinson’s [7, 8]. Therefore,
the present study was aimed at investigating the ef-
fects of coffee on body weight and serum biomarkers Experimental animals and protocol
of metabolic syndromes such as glucose, uric acid A total of 30 male albino Wistar rats of 6–7 weeks
and lipid profile levels in high fructose diet feeding were obtained from Addis Ababa University, depart-
male albino Wistar rats. ment of Pharmacology. The experimental rats were
placed in a plastic cage with stainless steel cover (5
rats/cage) and housed in biochemistry animal labora-
Methods and materials
tory with optimum temperature (24 + 1 °C), relative
Study area
humidity, optimum ventilation and 12-h light-dark
The study was carried out at Biochemistry Department La-
cycle. They were standardized for their feeding behav-
boratory, Addis Ababa University, Addis Ababa, Ethiopia.
ior and observed continuously for three weeks. Until
the initiation of the experiment, all rats were provided
Study period and duration
with free access to standard chow and plain tap
The study was undertaken for a period of 6 months from water. When the rats became 9–10 weeks old (adult),
July 2017 to January 2018. they were randomly assigned into six groups. Each rat
in the given group was identified by giving a number
Study design on its tail by permanent marker. Prior to the initi-
A post-test experimental study was conducted on rat ation of experiment, their body weight was measured
model in order to investigate the effect coffee on body by triple beam balance and ranged from 205 g to 245
weight, fasting serum glucose, uric acid and lipid profile g, with mean value of 225.4 ± 11.5 g. Then the rats
levels. were treated as follows:

Ethical consideration Group I (Normal control group): fed on standard

The ethical clearance was obtained from Department of chow and plain tap water only.
Biochemistry Research and Ethical Review Committee Group II (Fructose control group): fed on standard
(DRERC), by approval letter with Ref. No. of SOM/ chow and 20% (w/v) of fructose solution.
BCHM/154/2009, meeting No. of DRERC 02/17 and Group III (Treatment group): fed on standard chow
protocol No. of M.Sc. 11/17 issued on March 2/2017. and 20% (w/v) of fructose solution, and treated with 71
All experimental activities were carried out in accord- mg/kg BW/day of coffee.
ance with recommendations from the declaration of na- Group IV (Treatment group): fed on standard chow
tionally and internationally conventional standards for and 20% (w/v) of fructose solution, and treated with
the employment of experimental animals, and code of 142 mg/kg BW/day of coffee.
ethics of animal experiments, which comply with scien- Group V (Treatment group): fed on standard chow
tific and ethical guidelines. and 20% (w/v) of fructose solution, and treated with
213 mg/kg BW/day of coffee.
Study variables Group VI (Treatment group): fed on standard chow
Independent variable was coffee administration. Dependent and 20% (w/v) of fructose solution, and treated with
variables were body weight, Serum glucose, uric acid and 284 mg/kg BW/day of coffee.
lipid profile levels.
Preparation of fructose solution and coffee
Sample size determination Preparation of fructose solution
Sample size determination was based on WHO stand- Fructose used in this experiment was pure crystalline,
ard, which recommends that each treatment group of SIGMA-ALDRICH, USA, purchased from India.
experimental animal should be at least 5 animals [9]. Twenty percent (20%, W/V) of fructose solution [10]
The study was conducted on 6 groups of rats. was prepared on daily basis and substituted for tap
Feyisa et al. Laboratory Animal Research (2019) 35:29 Page 3 of 8

water in five groups of the rats (group II to group Blood sample collection
VI).The net consumption of each group was recorded At the end of the sixth week, the rats were fasted
in mL/day. overnight by removing fructose solution and and
standard chow, however, tap water was supplied for
Preparation of coffee and dosage calculation all groups of the rats. After overnight fasting the rats
Coffee used as a treatment in the present study was were anesthetized with diethyl ether. Blood sample
Coffea arabica (TO.MO.CA Coffee packet), pur- was collected by cardiac puncture and the rats were
chased from coffee shop in Addis Ababa. killed by exsanguination. To prepare serum, the blood
TO.MO.CA stands for Torrefazione Moderna Cafe sample was transferred into serum separator tube
(Italian), translated directly as modern coffee roast- (SST) and left to clot at room temperature for 30 min
ing. Taking into account the coffee brewing process immediately following collection. Subsequently, the
in Ethiopia, boiled, unfiltered coffee was prepared clotted blood sample was centrifuged at 2000 rpm for
following the instruction written on the packet (by 15 min. Finally, the serum was transferred into necked
adding 10 g of coffee powder into 180 mL of hot tube and stored at − 80 °C [12] until the analyses were
water. To estimate the amount of dissolved coffee performed.
powder, indirect method of measurement was used
(the residue of the coffee powder was measured and Laboratory tests
subtracted from the total added powder). Accord- Serum glucose, uric acid, TC, HDL-C and TG were de-
ingly, the volume of the coffee solution after boiling termined by an enzymatic colorimetric methods using
and decantation was 120 mL. The amount of undis- fully automated analyzer (Mindary BS-200E); LDL-C was
solved air-dried residue of coffee powder was 5.2 g, so calculated using Frieldwald’s formula.
the dissolved amount was calculated as 10 g–5.2 g =
4.8 g. The volume of 1 cup of coffee was as recently Data entry and analysis
used by Lelyana and Lelyana et al. (0.36 mL/200 g All data were entered and cleared by epi-data software
body weight/day) in rats, which is equivalent 125 mL/ version 3.1 and exported to SPSS (statistical Package
70 kg body weight/day of coffee in humans [9, 11]. In for Social Science) software version 23.0 for statistical
our study, the average body weight (BW) of the ex- analysis. Normality distributions were assessed by
perimental rats was 225 g. Shapiro-Wilks test and plots (stem-and-leaf and histo-
Accordingly, 16 mg/225 g BW/day of coffee was gram).One-way analysis of variance (ANOVA) was
considered as a single dose of coffee. Similarly, 0.8, done to determine statistical differences among all
1.2 and 1.6 mL/225 g BW/day (32 mg, 48 mg and 64 groups of the study. Pairwise comparisons were con-
mg/225 g BW/day) were considered as a double, triple ducted by Tukey post hoc multiple comparison
and quadruple doses of coffee. For the seek of con- tests.The results of the data were presented as mean ±
venience, these doses were converted to standard unit standard deviation (SD). The p-values < 0.05 were con-
(mg/kg) as 71, 142, 213 and 284 mg/kg BW/day sidered statistically significant.
respectively.
The total volume of coffee administered to treat- Results
ment groups and clear warm water to normal and Effects of coffee on energy intake
fructose control group by oral gavage was 2 mL. The The rats treated with 284 mg/kg BW/day of coffee
oral gavage was performed by the principal investiga- consumed lower amount of chow compared to fruc-
tor between 09:00 and 10:00 a.m. every day. The ex- tose control group (57.5 g/day vs 68.5 g/day) and nor-
periment was conducted for a period of 6 weeks. No mal control (57.5 g/day vs 95.5 g/day).The results also
rat died as a result of the treatment or other causes shows that the mean liquid intake in rats treated with
throughout the experiment. 284 mg/kg BW/day of coffee was lower when com-
pared to fructose control group (174.5 mL/day vs 187
Data collection mL/day). Since the chow and fructose solution were
Measurement of body weight supplied by treatment group, no statistical test was
Body weight of the experimental rats was measured by done (Fig. 1).
triple beam balance capable of measuring 610 ± 0.1 g at
initial and weekly during the experiment and recorded Effect of coffee on body weight
to the corresponding code of each rat in the group. Initially, the body weight of the rats was statistically
However, due to the fluctuation of body weight between similar among all groups (p = 0.96) (Fig. 2).
weeks, only the initial and final body weights were con- At end of the sixth week, the body weights of rats
sidered for final statistical test. treated with 213 and 284 mg/kg BW/day of coffee
Feyisa et al. Laboratory Animal Research (2019) 35:29 Page 4 of 8

Fig. 1 The mean consumption of chow and liquid of the rats. The values are expressed as mean. Sample size (n) is 5 for each group. Liquid -
refers to fructose solution for all groups, except for group I where it refers to plain tap water. Group I – Normal control group; Group II-Fructose
control group; Group (III- VI) –Treatment groups (received 71, 142, 213 and 284 mg/kg BW/day of coffee) respectively

Fig. 2 Initial body weight of the rats. Values are expressed as mean ± SD. Sample size (n) is 5 for each group I- Normal control group; II-Fructose
control group; III-VI -Treatment groups (received 71, 142, 213 and 284 mg/kg BW/day of coffee) respectively
Feyisa et al. Laboratory Animal Research (2019) 35:29 Page 5 of 8

Fig. 3 Final body weight of the rats. Values are expressed as mean ± SD. Sample size (n) is 5 for each group I- Normal control group; II-Fructose
control group; III-VI -Treatment groups (received 71, 142, 213 and 284 mg/kg BW/day of coffee) respectively

significantly reduced compared to fructose control to normal control (p = 0.007, 0.002) respectively
group (p = 0.047; 0.035) respectively. In addition, body (Table 1).
weight of fructose control group was significantly
higher compared to normal control group (p = 0.020) Effect of coffee on serum lipid profile levels
(Fig. 3). Fasting serum TG and LDL-C levels were significantly
lower in rats treated with 284 mg/kg BW/day of coffee
compared to fructose control group (p = 0.031; 0.046) re-
Effect of coffee on serum glucose and uric acid levels spectively. Similarly, significant elevation of fasting
Treating the rats with 213 and 284 mg/kg BW/day of serum TG (p = 0.013) and LDL-C (p = 0.007) levels were
coffee significantly decreased fasting serum glucose also found in fructose control group when compared to
(p = 0.049; 0.029) and uric acid (p = 0.026; 0.010) normal control group. Significantly increased fasting
levels compared to fructose control group. Fasting serum LDL-C level was also observed in rats treated
serum glucose and uric acid levels were significantly with 71 mg/kg BW/day of coffee as compared to normal
increased in fructose control group when compared control group (p = 0.046), (Table 2).

Table 1 Fasting serum glucose and uric acid levels of the rats
Variable Groups F P
(mg/dL)
I II III IV V VI
Glucose 74.2 ± 8 94.4 ± 9.6abc 88.8 ± 9 81 ± 5.9 78.4 ± 8.4 77.2 ± 7.5 4.5 0.005*
abc
Uric acid 1.32 ± 0.3 2.18 ± 0.3 1.7 ± 0.4 1.64 ± 0.3 1.52 ± 0.26 1.44 ± 0.3 4.7 0.004*
*-Indicates significant differences among all groups at p < 0.05 as tested by one-way ANOVA. Superscript letters (a, b and c) indicate significant differences
compared to group I, V and VI respectively at p < 0.05 as tested by Tukey post hoc multiple comparisons
 Feyisa et al. Laboratory Animal Research (2019) 35:29 Page 6 of 8

Table 2 Fasting serum lipid profile levels of the experimental rats

Variable Groups FP
(mg/dL)
I II III IV V VI
TC 101.4 ± 13.2 120.2 ± 12.4 118.6 ± 8.8 113.2 ± 10.2 106.6 ± 9.4 103.8 ± 10.7 2.6 0.051
HDL-C 44 ± 5.5 33.4 ± 5.5 37.6 ± 4.5 39 ± 4.3 39.8 ± 6.8 41 ± 7.3 2 0.132
TG 116.8 ± 11 147.8 ± 17.8ab 142.6 ± 13.2 135 ± 15.4 125.4 ± 10.6 120 ± 9.8 4.5 0.005*
ab a
LDL-C 34 ± 9.5 57.2 ± 10.4 52.5 ± 4.4 47.2 ± 13.5 41.7 ± 6.7 38.8 ± 8.8 4.4 0.006*
HDL-C- High density lipoprotein cholesterol; LDL-C- Low density lipoprotein cholesterol; TC- Total cholesterol; TG- Triglycerides. Sample size (n) is 5 for each
group.* -Indicates significant difference among all groups at p < 0.05 as tested by one-way ANOVA. Superscript letters (a) - indicates significant differences
compared to group I; (b) - indicates significant differences compared to group VI at p < 0.05 as tested by Tukey post hoc multiple comparisons

Discussion Most probably, coffee prevented increase in glucose

The obtained results indicated that significantly de- level due to its main component, caffeine, which inhibits
creased body weight gain was recorded in rats treated adenosine receptors that stimulates hepatic glucose pro-
with 213 and 284 mg/kg BW/day of coffee at the end of duction through the activation of A2B adenosine recep-
sixth week as compared to fructose control group (p = tors [21]. Caffeine might also have stimulated glucose
0.047; 0.035) respectively. These results suggest that the transport through activation of cyclic AMP-dependent
effect of coffee on body weight is dose dependent. In line protein kinase α-1 [20]. Another mechanism could be
with this finding, Ismail et al. reported that male albino via selective inhibition of hepatic glucose-6-phosphatase,
rats fed on different preparations of Turkish, Arabian a rate-limiting enzyme of gluconeogenesis, by CGA of
and instant coffee had significantly smaller weight gain coffee [3, 22].
than normal control [13]. Similar finding was also re- Fasting serum glucose was significantly higher in
ported by Mohmoud et al. in which feeding on different fructose control group than in normal control group
doses of Arabic coffee for 30 days lowered body weight (p = 0.007). This finding is in agreement with Sandeva
in rats fed on basal diet [14]. The possible mechanism et al.’s study, which reported that serum glucose level
by which coffee prevented higher body weight gain in was significantly increased in male and female rats
done the present study could be by increasing lipolysis via cat- fed on 20% of fructose solution in drinking water for
echolamines [15, 16]. Caffeine might also have caused 8 weeks [23].
body weight loss by increasing physical activity. It was found that fasting serum uric acid levels were
In addition, coffee had decreased energy intake, which significantly lower in rats treated with 213 and 284 mg/
might in turn had led to decreased in the body weight kg BW/day of coffee compared to fructose control group
gain. In comparison with normal control group, fructose (p = 0.026; 0.010) respectively. Similar studies by Lelyana
control group had statistically significant body weight and Lelyanaet al. reported that coffee had non-
gain (p = 0.020). This result is in line with Tanaka et al.’s significantly reduced serum uric acid in obese rats and
report in which feeding on 20% fructose solution in tap mice with hyperuricemia respectively [9, 11]. In addition,
water for 8 weeks had brought statistically significant Mahmoud et al. reported that different doses of Arabic
body weight gain in both sexes of rats [1]. As demon- coffee had lowered serum uric acid levels in experimen-
strated in both short-term and long-term studies, fruc- tal rats fed on basal diet [14]. In comparison with men-
tose consumption results in decreased circulating levels tioned findings, the significant decrease in fasting serum
of insulin and leptin and increased caloric intake [17]. uric acid found in the present study might be due to
Treating rats with 213 and 284 mg/kg BW/day of cof- variation in the doses of coffee, study design and/or
fee had significantly reduced fasting serum glucose levels duration. Coffee consumption may lower serum uric
as compared to fructose control group (p = 0.049; 0.029) acid levels and risk of gout via various mechanisms.
respectively. Consistent with these findings, Li et al. re- The main mechanism is via competitive inhibition of
ported that CGA significantly lowered fasting glucose xanthine oxidase by caffeine (1,3,7-trimethyl-
levels in golden hamsters [18]. In addition, Takami et al. xanthine), [24]. The other mechanism is probably by a
reported moderate coffee consumption in human sub- diuretic action of caffeine that might affect serum uric
jects was associated with a significantly lower odds ratio acid concentration through increased excretion in
for high plasma glucose [19].The overall effects of coffee urine [21].
on glucose level in the present study revealed good im- Evaluation of fasting serum lipid profiles showed that
plication to support the hypothesis that states, “Heavy treating rats with 284 mg/kg BW/day of coffee had signifi-
coffee consumption has been associated with a lower cantly lowered fasting serum levels of TG and LDL-C
risk of diabetes,” [20]. when compared to fructose control group (p = 0.031;
Feyisa et al. Laboratory Animal Research (2019) 35:29 Page 7 of 8

0.046) respectively. Consistent with these findings, Gomes findings of the present study suggest that coffee con-
et al. reported that feeding of 7.2 mL/kg of body weight sumption may be helpful in ameliorating metabolic syn-
for 41 days significantly lowered lipid percentage in hyper- dromes and its associated complications such as obesity,
lipidemic diet feeding rats [25]. In addition, Li et al. re- diabetes, inflammation and cardiovascular diseases.
ported CGA significantly lowered the levels of fasting
Abbreviations
serum TG, FFA, TC, and LDL-C in golden hamsters [18]. ANOVA: Analysis of Variance; BW: Body Weight; cAMP: Cyclic Adenine
Fasting serum levels of TC and HDL-C were non- Monophosphate; CGA: Chlorogenic Acid; FFA: Free Fatty Acid; gm: Gram;
significantly lower and higher respectively, in rats HDL-C: High Density Lipoprotein Cholesterol; Kg: Kilogram; LDL-C: Low
Density Lipoprotein Cholesterol; mg/dL: Milligram per deciliter;
treated with 213and 284 mg/kg BW/day of coffee com- rpm: Revolution per Minute; SD: Standard Deviation; T2DM: Type 2 Diabetes
pared to fructose control group (Table 3).These results Mellitus; TC: Total Cholesterol; TG: Triglycerides; TL: Total Lipid; UA: Uric Acid;
are in agreement with Karabudak et al.’s finding that W/V: Weight Per Volume
reported Turkish and instant coffee consumption did Acknowledgements
not significantly affect serum levels of TC among Turk- We would like to express our deepest gratitude to Haramaya University and
ish subjects [26]. On the other hand, in Mohmoud Addis Ababa for funding this research project. We are extremely grateful to the
Department of Medical Biochemistry, Addis Ababa University, for its
et al.’s study,Arabic coffee had significantly decreased indispensable support and coordination in smooth running of this project. We
serum TC levels in experimental rats fed on basal diet would also like to express our special appreciation to Mr. Abdissa Tufa for his
[14]. In contradiction to our findings, few studies re- indispensable comments and suggestions and Mrs. Ture Girma for her active
cooperation and contribution during the entire experiment of the study.
ported significant elevation of serum TL, TC, TG and
LDL-C, while a significant decrease of HDL-C in rats Authors’ contributions
fed diet supplemented with low or high dose of coffee. TOF conceived and designed the study, conducted the experiment,
The causes for these variations are unclear. However, it collected, analyzed and interpreted the data, and wrote the manuscript. DSM
designed the study, interpreted the data and advised me, MM and WLL
may be due to the variations in the mode of coffee admin- advised me. MLH assisted in data data analysis and writing the manuscript.
istration. Lipid lowering effects of coffee found in the All the authors read, commented on, and contributed to the submitted and
present study could be related to caffeine, (methylxan- revised manuscript. All authors read and approved the final manuscript.

thine) which stimulates lipolysis by increasing cellular Funding

levels of cAMP through antagonism adenosine receptors The research was funded by Haramaya University and Addis Ababa
(A1) and inhibition of phosphodiesterase activity [25, 27]. University. The funders had no role in designing of the study, collection,
analysis and interpretation of data as well as in the writing of the
Feeding rats on 20% of fructose solution for six weeks manuscript.
significantly increased fasting serum levels of TG in fruc-
tose control (p = 0.013); LDL-C in fructose control (p = Availability of data and materials
All necessary data and materials related to the article are included in the
0.007) and 71 mg/kg BW/day of coffee treated(p = 0.046) manuscript.
groups compared to normal control group.The fasting
serum levels of TC (p = 0.106) and HDL-C (p = 0.073) Competing interests
The authors declare that they have no competing interests.
were non-significantly higher and lower,respectively in
fructose control group than in normal control group Author details
1
(Table 3).Fasting serum levels of TC, TG, and LDL-C Department of Medical Biochemistry, College of Health and Medical
Sciences, Haramaya University, Harar, Ethiopia. 2College of Health Sciences,
were possibly increased in fructose control group as me-
Addis Ababa University, Addis Ababa, Ethiopia.
tabolism of fructose bypasses the enzyme phosphofruc-
tokinase and unregulated the flow of fructose-derived Received: 31 July 2019 Accepted: 5 November 2019
substrates (glycerol-3- phosphate and acetyl-CoA) into
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