research article mechanism of body weight reducing effect of...

Hindawi Publishing CorporationInternational Journal of EndocrinologyVolume 2013 Article ID 914651 5 pageshttpdxdoiorg1011552013914651

Research ArticleMechanism of Body Weight Reducing Effect ofOral Boric Acid Intake

Erhan Aysan1 Fikrettin Sahin2 Dilek Telci2 Merve Erdem2 Mahmut Muslumanoglu1

Erkan YardJmcJ1 and Huseyin Bektasoglu1

1 Bezmialem Vakif University ATA-2 Sitesi Akasya Caddesi No 25 Cengelkoy Uskudar 80700 Istanbul Turkey2Department of Genetics and Bioengineering Yeditepe University Turkey

Correspondence should be addressed to Erhan Aysan erhanaysanhotmailcom

Received 1 March 2013 Accepted 14 May 2013

Academic Editor Oreste Gualillo

Copyright copy 2013 Erhan Aysan et al This is an open access article distributed under the Creative Commons Attribution Licensewhich permits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

Objective The effect of oral boric acid intake on reducing body weight has been previously demonstrated although the mechanismhas been unclear This research study reveals the mechanism Subjects Twelve mice were used in groups of six each in the controland study groups For five days control group mice drank standard tap water while during the same time period the study groupmice drank tapwaterwhich contains 028mg250mLboric acid After a 5-day period gene expression levels for uncoupling proteins(UCPs) in the white adipose tissue (WAT) brown adipose tissue (BAT) and skeletal muscle tissue (SMT) and total body weightchanges were analyzed Results Real time PCR analysis revealed no significant change in UCP3 expressions but UCP2 inWAT (11987500317) BAT (119875 0014) and SMT (119875 00159) and UCP1 in BAT (119875 0026) were overexpressed in the boric acid group In additionmice in the boric acid group lost body weight (mean 281) while mice in the control group experienced no weight loss but a slightweight gain (mean 009 119875 lt 0001) Conclusion Oral boric acid intake causes overexpression of thermogenic proteins in theadipose and skeletal muscle tissues Increasing thermogenesis through UCP protein pathway results in the accelerated lipolysis andbody weight loss

1 Introduction

Boron immediately to the left of carbon atom in the periodictable is so similar to carbon thatmany carbon-basedmolecu-les are for all practical purposes the same as boron-basedmolecules Boron is a stable metal found in nature as borateThe actual mean daily intake of boron in human diet is esti-mated to be 12mgday [1] Boron is used in a wide range ofproducts including glass detergents fire retardants andfibers to reinforce the plane fuselages and body armor and inother superhard materials [2] The similarity of boron to car-bon has led to its wide use in biology [1 3ndash5]

Recently we demonstrated that oral boric acid intakelowered the body weight of BALBc outbred female mice [6]Its mechanism however remained unclear In this researchwe aim to reveal the molecular mechanism underlying thebody weight reducing the effect of oral boric acid intake Todo so fresh white adipose tissue (WAT) brown adipose tissue(BAT) and skeletal muscle tissue (SMT) were dissected from

mice in the control group without the boric acid intake andthe experimental group with the boric acid intake and weanalyzed for the changes in the expression levels of uncou-pling proteins (UCPs) 1 2 and 3 using the real time PCRanalysis

UCPs are transmembrane proteins in the inner mem-branes ofmitochondria and are responsible for themitochon-drial proton leak Proton transport through UCPs lowers themitochondrial membrane potential leaving fewer protonsto be transferred down the electrochemical gradient by theF0F1 ATP synthase

The proton electrochemical gradient is dissipated as heatrather than being converted into ATP resulting in the mobi-lization of fatty acid stocks without a consequent increase inATP production [7] Emerging evidence has suggested thatdifferences in UCP expression among individuals may beresponsible for the wide range of metabolic rate among pop-ulations Reduced levels of UCPmRNA were associated withthe increased risks for obesity in mice and humans [8ndash10]

2 International Journal of Endocrinology

In this paper we demonstrate for the first time that theoral administration of boric acid induces an increase in theexpression levels of UCP2 in WAT BAT and SMT cells alsoUCP1 in BAT cells

2 Methods

This study was performed in the Bezmialem Vakif UniversityExperimental Animals Research Laboratory and YeditepeUniversity Genetics and Bioengineering Department Labo-ratories Research protocol was approved by the BezmialemVakif University Local Animal Ethics Committee All steps inthis research were in accordance with the regulations govern-ing the care and use of laboratory animals as set forth in theDeclaration of Helsinki

Twelve outbred produced 8-week-old female BALBcmice were divided into two groups According to poweranalysis with 005 accuracy and 095 power the number ofmice was determined as 6 each in both the control and studygroups The animals were kept in standard metabolic cagesdesigned specifically for mice A daily cycle of 12 hours oflightand 12 hours of dark was used for illumination of theroom where mice were placed

For five days mice were fed ad libitum with standardpellet feed manufactured specifically for small animals Thestudy group mice ingested boric acid through their drinkingwater 028mg boric acid (2mg Bor Atac DF 14 Boric AcidTMT Co Tekirdag Turkey) was added to 250mL tap waterand dissolved by 3 minutes of agitation

On days 0 through 5 all the mice were weighed everyafternoon and the data were recorded After five days the ani-mals were sacrificed by means of cervical dislocation

WATs were isolated from omentum and SMTs were iso-lated from anterior abdominal muscles via anterior midlineabdominal incision BATs were isolated from interscapulararea via posterior midline thoracocervical incision

Tissues were evaluated for changes in the expressionlevels of UCPs 1 2 and 3 using the real time polymerase chainreaction (PCR) analysis The primary evaluation parameterof this research was analysis of alterations of gene expressionlevels for UCP1 UCP2 and UCP3 in WAT BAT and SMTcells The secondary evaluation parameter was total bodyweight change observed in both groups of mice

3 RNA Isolation and Real Time PCR Analysis

Total RNA from the fresh WAT BAT and SMT was iso-lated using peqGOLD RNAPure reagent (Peqlab Germany)according to themanufacturerrsquos instructions Quality of RNAwas checked by spectrophotometry and agarose gel elec-trophoresis cDNA synthesis was performed using RevertAidFirst Strand cDNA Synthesis Kit (Fermentas Lith-uania) at42∘C for 60min and 70∘C for 5min as described by themanufacturer Real time PCR was performed with Quan-tiTect SYBR green PCR kit (Qiagen Germany) accordingto instructions using 03120583M primer concentration PCRincluded 94∘C for 15min initial denaturation step followedby 40 cycles of 94∘C for 30 sec annealing for 1min (61∘C for

UCP1 and 18S rRNA 58∘C for UCP2 55∘C for UCP3) and72∘C for 1min and a final extension step 72∘C for 10minEach sample was analyzed in triplicate and the speci-ficity of products was checked by the melt curve analysisResults were analyzed by the standard curve method usingthe 18S rRNA house-keeping gene for normalization Thesequences of the primers were as follows for 18S rRNAsense 51015840-AACTGAGGCCATGATTAAGAGG-31015840 antisense51015840-GGCATCGTTTATGGTTGGAAC-31015840 for UCP1 sense51015840-CTCGGGTCCTGGAACGTCAT-31015840 antisense 51015840-CAA-CGGAGCTGTTCATTTGATTTC-31015840 for UCP2 sense 51015840-GCTGGTGGTGGTCGGAGATA-31015840 antisense 51015840-ACA-GTTGACAATGGCATTACGG-31015840 [10] and for UCP3 sense51015840-GGCTGCCTGGAACAGAACAA-31015840 and antisense 51015840-TCCCATCAGGTCAGTGCAAAAC-31015840 All real time PCRexper-iments were performed using iCycler iQ real time PCRdetection system (Bio-Rad USA)

4 Statistical Analysis

Statistical analyses were performed using SigmaStat SoftwareResults were evaluated with a confidence interval of 95 anda 119875 lt 005 level In addition to descriptive statistical methods(mean standard deviation and median) the Mann-Whitney119880 test was used for the inter-group comparisons and theWilcoxon test for comparison of in-group variables

5 Results

Real time PCR analyses revealed that boric acid intakeincreased the UCP2 expression inWAT by 27-fold (119875 00317Figure 1) BAT by 105-fold (119875 0014 Figure 2) and SMT by45-fold (119875 00159 Figure 3) (Table 1) In addition boric acidintake led to a 9-fold (119875 0026) increase in UCP1 expressionin BAT (Figure 2) UCP1 in the other two tissues andUCP3 inall three tissues were not overexpressed significantly (119875 gt005 Figure 1 Table 1)

Bodyweight changes of the groups and statistical analyseswere revealed in Table 2 Mice which were fed with boric acidlost their body weight at a mean of 281 in five days In thecontrol group no weight loss was observed indeed a weightgain averaged at 009 was recorded Body weight changesbetween the groups were statistically significant (119875 lt 0001)From day 0 through day 5 total body weight differences werestatistically significant in the boric acid group (119875 lt 0005)but not in the control group

6 Discussion

It is clear that obesity is characterized by an imbalancebetween energy intake and expenditure Emerging evidencehas placed adipocytes at the center stage since alterationsin the regulation of lipogenesis and lipolysis have beenassociated with obesity Oberkofler et al demonstrated thatimpaired adipose tissue expression ofUCP2may play a role inthe pathophysiology of obesity [10]

UCP1 is a key protein in thermogenesis and regulationof energy expenditure mechanisms which are important in

International Journal of Endocrinology 3

ControlBoric acid

times10minus4

012

010

008

006

004

002

0

P = 00317

UCP1 UCP2 UCP3

mRN

A ex

pres

sion

18Sr

RNA

Figure 1 Real time PCR analyses in WAT (119875 00317)

obesity [11] Molecular players that are responsible for thecontrol of UCP expression in adipocytes are still a subjectof debate however hormonal stimuli such as glucagonand triiodothyronine treatment or feedingfasting conditionshave been demonstrated to affect the UCP-2 expression inadipocytes [7]

In our first-step research as discussed above we hadrevealed the effect of oral boric acid intake on reducingbody weight [6] In our previous research we revealed thatblood cholesterol LDL AST ALT LDH amylase and urineurobilinogen levels were statistically high in the boric acidintake group and therefore we hypothesized that these resultsmay be related to an increased catabolismwith especially highlipid consumptionNow in the second step of our researchwehave revealed the mechanism the boric acid intake led to theover expression of UCPs (UCP1 in BAT UCP2 in WAT BATand SMT) resulting in an increase in thermogenesis and anacceleration of lipolysis

The function of adipocytes is the storage of triacylglyc-erols as energy reserve in the body and maintenance of thesystemic energy balance through the regulation of lipogenesisand lipolysis In addition adipocytes such as the ones foundin the brown fat tissue are involved in the thermoregulatorythermogenesis where the energy from the cellular respirationis transferred into heat by the action of the mitochondrialuncoupling proteins (UCP-1) UCP-1 the first UCP isolatedand identified is highly expressed in the brown fat tissue[12] and involved in uncoupling of proton transport fromATP synthesis hence increase the demand for fuel substratessuch as glucose and lipids in the oxidative phosphorylationprocesses [13] Additions to the UCP family came with thediscovery of the homologues UCP2 and UCP3 [14 15] WhileUCP2 was found to be widely expressed in different rangeof cells UCP3 expression was mainly restricted to skeletaland cardiac muscle and brown fat [7] Although the contri-bution of UCP2 and UCP3 in nonshivering thermogenesis isdebatable due to their low expression ratios in themembranerecent reports suggested that UCP2 and UCP3 similar to

ControlBoric acid

times10minus4

25

20

15

10

5

0

P = 0026

P = 0014

UCP1 UCP2 UCP3

mRN

A ex

pres

sion

18Sr

RNA

Figure 2 Real time PCR analyses in BAT (119875 0014)

P = 00159

ControlBoric acid

UCP1 UCP2 UCP3

times10minus4

2

15

1

05

0

mRN

A ex

pres

sion

18Sr

RNA

Figure 3 Real time PCR analyses in SMT (119875 00159)

UCP1 can lower the proton electrochemical gradient acrossthe mitochondrial membrane causing a decrease in ATP pro-duction when overexpressed in mammalian cells [13 16]

Although effects of boron on metabolism and enzymeswere known its body weight reducing effect through UCP1and UCP2 overexpression was not demonstrated before It isclear that obesity is characterized by an imbalance betweenenergy intake and expenditure Emerging evidence hasplaced adipocytes at center stage since alterations in theregulation of lipogenesis and lipolysis have been associatedwith obesity For example isolated adipocytes from obesepatients showed a twofold decrease in the specific activityof the lipogenicmarker enzymeGlyceraldehyde-3-PhosphateDehydrogenase (G3PDH) when compared to that of nono-bese patients resulting in a lower basal lipolytic rate [17]There is also recent evidence relating the reduced UPC2expression to morbid obesity hence increasing the emphasison the importance of adipocytes in maintaining homeostasis


Table 1 The mean gene expression levels normalized against 18sRNA plusmn SE expression for each UCP proteins in WAT BAT and SMT

Boric acid group Control group P valueUCP1 mRNA in WAT 000000017 plusmn 000000004 000000015 plusmn 000000008 06905UCP2 mRNA in WAT 000000527 plusmn 000000420 000000019 plusmn 000000006 00317UCP3 mRNA in WAT 000000013 plusmn 000000007 000000012 plusmn 000000006 09452UCP1 mRNA in BAT 000025809 plusmn 000011527 000002859 plusmn 000001483 0026UCP2 mRNA in BAT 000123186 plusmn 000067987 000011691 plusmn 000006580 0014UCP3 mRNA in BAT 000002918 plusmn 000000762 000001676 plusmn 000000534 02949UCP1 mRNA in SMT 000000071 plusmn 000000022 000000084 plusmn 000000063 0366UCP2 mRNA in SMT 000002884 plusmn 000000967 000000645 plusmn 000000198 00159UCP3 mRNA in SMT 000011747 plusmn 000005621 000009499 plusmn 000003876 07308

Table 2 Body weight changes of the groups and statistical analysesin five days

Groups Day 0 Day 5 Difference Wilcoxon Test119911 P

Control GroupMean 2150 2192 042 minus1890 0059Std Deviation 105 132 038

Boric Acid GroupMean 2183 1592 minus592 minus2220 0026Std Deviation 117 080 066

Mann-WhitneyTest119911 minus580 minus2892 minus2929

P 562 004 003

of energy dissipation and storage [9] Molecular playersthat are responsible for the control of UCP expression inadipocytes are still a subject of debate However hormonalstimuli such as glucagon and triiodothyronine treatment orfeedingfasting conditions have been demonstrated to affectthe UCP-2 expression in adipocytes [7]

In conclusion our results are promising and may open anew line inquiry for the boron-based prevention or treatmentapproaches to obesity Some questions await answers suchas complications of long-term boric acid intake The effect ofboric acid intake must also be evaluated as to different means(local muscular or intravenous) and with different doses AsUCPs may not be as active in other animals such as rat andrabbit as it was in mice their effect should also be tested inthese model animals Taken together boric acid intake maybecome a new and effective way to treat the worldwide healthproblem of obesity

Acknowledgment

The authors would like to thank S Delacroix for English edit-ing of this paper

References

[1] D L AndersonWC Cunningham andT R Lindstrom ldquoCon-centrations and intakes of H B S K Na Cl andNaCl in foodsrdquo

Journal of Food Composition and Analysis vol 7 no 1-2 pp 59ndash82 1994

[2] P Hunter ldquoNot boring at all Boron is the new carbon in thequest for novel drug candidatesrdquo EMBO Reports vol 10 no 2pp 125ndash128 2009

[3] S Altieri S Bortolussi P Bruschi et al ldquoNeutron autoradiog-raphy imaging of selective boron uptake in human metastatictumoursrdquo Applied Radiation and Isotopes vol 66 no 12 pp1850ndash1855 2008

[4] K Warington ldquoThe effect of boric acid and borax on the broadbean and certain other plantsrdquo Annals of Botany vol 37 no 4pp 629ndash672 1923

[5] A F Rossi R DMiles B L Damron and L K Flunker ldquoEffectsof dietary boron supplementation on broilersrdquo Poultry sciencevol 72 no 11 pp 2124ndash2130 1993

[6] E Aysan F Sahin D Telci et al ldquoBodyweight reducing effect oforal boric acid intakerdquo International Journal of Medical Sciencesvol 8 pp 653ndash658 2011

[7] J Mozo Y Emre F Bouillaud D Ricquier and F CriscuololdquoThermoregulation what role for UCPs in mammals andbirdsrdquo Bioscience Reports vol 25 no 3-4 pp 227ndash249 2005

[8] J Kopecky G Clarke S Enerback B Spiegelman and L PKozak ldquoExpression of the mitochondrial uncoupling proteingene from the aP2 gene promoter prevents genetic obesityrdquoJournal of Clinical Investigation vol 96 no 6 pp 2914ndash29231995

[9] H Oberkofler G Dallinger Y M Liu E Hell F Krempler andW Patsch ldquoUncoupling protein gene quantification of expres-sion levels in adipose tissues of obese and non-obese humansrdquoJournal of Lipid Research vol 38 no 10 pp 2125ndash2133 1997

[10] H Oberkofler Y M Liu H Esterbauer E Hell F Kremplerand W Patsch ldquoUncoupling protein-2 gene reduced mRNAexpression in intraperitoneal adipose tissue of obese humansrdquoDiabetologia vol 41 no 8 pp 940ndash946 1998

[11] H M Feldmann V Golozoubova B Cannon and J Neder-gaard ldquoUCP1 ablation induces obesity and abolishes diet-induced thermogenesis in mice exempt from thermal stress byliving at thermoneutralityrdquo Cell Metabolism vol 9 no 2 pp203ndash209 2009

[12] D Ricquier and J C Kader ldquoMitochondrial protein alterationin active brown fat a sodium dodecyl sulfate polyacrylamidegel electrophoretic studyrdquo Biochemical and Biophysical ResearchCommunications vol 73 no 3 pp 577ndash583 1976

[13] P Gonzalez-Muniesa F I Milagro J Campion and J AMartınez ldquoEctopic UCP1 gene expression inHepG2 cells affectsATP productionrdquo Journal of Physiology and Biochemistry vol61 no 2 pp 389ndash394 2005


[14] C Fleury M Neverova S Collins et al ldquoUncoupling protein-2 a novel gene linked to obesity and hyperinsulinemiardquo NatureGenetics vol 15 no 3 pp 269ndash272 1997

[15] O Boss S Samec A Paoloni-Giacobino et al ldquoUncouplingprotein-3 a new member of the mitochondrial carrier familywith tissue-specific expressionrdquo FEBS Letters vol 408 no 1 pp39ndash42 1997

[16] C Garcıa-Martinez B Sibille G Solanes et al ldquoOverexpressionofUCP3 in cultured humanmuscle lowersmitochondrialmem-brane potential raises ATPADP ratio and favors fatty acid vsglucose oxidationrdquoThe FASEB Journal vol 15 no 11 pp 2033ndash2035 2001

[17] B B Lowell and J S Flier ldquoBrown adipose tissue 1205733-adrenergicreceptors and obesityrdquo Annual Review of Medicine vol 48 pp307ndash316 1997

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Evidence-Based Complementary and Alternative Medicine

Volume 2014Hindawi Publishing Corporationhttpwwwhindawicom


In this paper we demonstrate for the first time that theoral administration of boric acid induces an increase in theexpression levels of UCP2 in WAT BAT and SMT cells alsoUCP1 in BAT cells

2 Methods

This study was performed in the Bezmialem Vakif UniversityExperimental Animals Research Laboratory and YeditepeUniversity Genetics and Bioengineering Department Labo-ratories Research protocol was approved by the BezmialemVakif University Local Animal Ethics Committee All steps inthis research were in accordance with the regulations govern-ing the care and use of laboratory animals as set forth in theDeclaration of Helsinki

Twelve outbred produced 8-week-old female BALBcmice were divided into two groups According to poweranalysis with 005 accuracy and 095 power the number ofmice was determined as 6 each in both the control and studygroups The animals were kept in standard metabolic cagesdesigned specifically for mice A daily cycle of 12 hours oflightand 12 hours of dark was used for illumination of theroom where mice were placed

For five days mice were fed ad libitum with standardpellet feed manufactured specifically for small animals Thestudy group mice ingested boric acid through their drinkingwater 028mg boric acid (2mg Bor Atac DF 14 Boric AcidTMT Co Tekirdag Turkey) was added to 250mL tap waterand dissolved by 3 minutes of agitation

On days 0 through 5 all the mice were weighed everyafternoon and the data were recorded After five days the ani-mals were sacrificed by means of cervical dislocation

WATs were isolated from omentum and SMTs were iso-lated from anterior abdominal muscles via anterior midlineabdominal incision BATs were isolated from interscapulararea via posterior midline thoracocervical incision

Tissues were evaluated for changes in the expressionlevels of UCPs 1 2 and 3 using the real time polymerase chainreaction (PCR) analysis The primary evaluation parameterof this research was analysis of alterations of gene expressionlevels for UCP1 UCP2 and UCP3 in WAT BAT and SMTcells The secondary evaluation parameter was total bodyweight change observed in both groups of mice

3 RNA Isolation and Real Time PCR Analysis

Total RNA from the fresh WAT BAT and SMT was iso-lated using peqGOLD RNAPure reagent (Peqlab Germany)according to themanufacturerrsquos instructions Quality of RNAwas checked by spectrophotometry and agarose gel elec-trophoresis cDNA synthesis was performed using RevertAidFirst Strand cDNA Synthesis Kit (Fermentas Lith-uania) at42∘C for 60min and 70∘C for 5min as described by themanufacturer Real time PCR was performed with Quan-tiTect SYBR green PCR kit (Qiagen Germany) accordingto instructions using 03120583M primer concentration PCRincluded 94∘C for 15min initial denaturation step followedby 40 cycles of 94∘C for 30 sec annealing for 1min (61∘C for

UCP1 and 18S rRNA 58∘C for UCP2 55∘C for UCP3) and72∘C for 1min and a final extension step 72∘C for 10minEach sample was analyzed in triplicate and the speci-ficity of products was checked by the melt curve analysisResults were analyzed by the standard curve method usingthe 18S rRNA house-keeping gene for normalization Thesequences of the primers were as follows for 18S rRNAsense 51015840-AACTGAGGCCATGATTAAGAGG-31015840 antisense51015840-GGCATCGTTTATGGTTGGAAC-31015840 for UCP1 sense51015840-CTCGGGTCCTGGAACGTCAT-31015840 antisense 51015840-CAA-CGGAGCTGTTCATTTGATTTC-31015840 for UCP2 sense 51015840-GCTGGTGGTGGTCGGAGATA-31015840 antisense 51015840-ACA-GTTGACAATGGCATTACGG-31015840 [10] and for UCP3 sense51015840-GGCTGCCTGGAACAGAACAA-31015840 and antisense 51015840-TCCCATCAGGTCAGTGCAAAAC-31015840 All real time PCRexper-iments were performed using iCycler iQ real time PCRdetection system (Bio-Rad USA)

4 Statistical Analysis

Statistical analyses were performed using SigmaStat SoftwareResults were evaluated with a confidence interval of 95 anda 119875 lt 005 level In addition to descriptive statistical methods(mean standard deviation and median) the Mann-Whitney119880 test was used for the inter-group comparisons and theWilcoxon test for comparison of in-group variables

5 Results

Real time PCR analyses revealed that boric acid intakeincreased the UCP2 expression inWAT by 27-fold (119875 00317Figure 1) BAT by 105-fold (119875 0014 Figure 2) and SMT by45-fold (119875 00159 Figure 3) (Table 1) In addition boric acidintake led to a 9-fold (119875 0026) increase in UCP1 expressionin BAT (Figure 2) UCP1 in the other two tissues andUCP3 inall three tissues were not overexpressed significantly (119875 gt005 Figure 1 Table 1)

Bodyweight changes of the groups and statistical analyseswere revealed in Table 2 Mice which were fed with boric acidlost their body weight at a mean of 281 in five days In thecontrol group no weight loss was observed indeed a weightgain averaged at 009 was recorded Body weight changesbetween the groups were statistically significant (119875 lt 0001)From day 0 through day 5 total body weight differences werestatistically significant in the boric acid group (119875 lt 0005)but not in the control group

6 Discussion

It is clear that obesity is characterized by an imbalancebetween energy intake and expenditure Emerging evidencehas placed adipocytes at the center stage since alterationsin the regulation of lipogenesis and lipolysis have beenassociated with obesity Oberkofler et al demonstrated thatimpaired adipose tissue expression ofUCP2may play a role inthe pathophysiology of obesity [10]

UCP1 is a key protein in thermogenesis and regulationof energy expenditure mechanisms which are important in


ControlBoric acid

times10minus4

012

010

008

006

004

002

0

P = 00317

UCP1 UCP2 UCP3

mRN

A ex

pres

sion

18Sr

RNA





ControlBoric acid

times10minus4

25

20

15

10

5

0

P = 0026

P = 0014

UCP1 UCP2 UCP3

mRN

A ex

pres

sion

18Sr

RNA


P = 00159

ControlBoric acid

UCP1 UCP2 UCP3

times10minus4

2

15

1

05

0

mRN

A ex

pres

sion

18Sr

RNA












P 562 004 003



Acknowledgment


References

























of






Disease Markers



OncologyJournal of





PPAR Research



Journal of

ObesityJournal of

















ControlBoric acid

times10minus4

012

010

008

006

004

002

0

P = 00317

UCP1 UCP2 UCP3

mRN

A ex

pres

sion

18Sr

RNA





ControlBoric acid

times10minus4

25

20

15

10

5

0

P = 0026

P = 0014

UCP1 UCP2 UCP3

mRN

A ex

pres

sion

18Sr

RNA


P = 00159

ControlBoric acid

UCP1 UCP2 UCP3

times10minus4

2

15

1

05

0

mRN

A ex

pres

sion

18Sr

RNA












P 562 004 003



Acknowledgment


References

























of






Disease Markers



OncologyJournal of





PPAR Research



Journal of

ObesityJournal of
























P 562 004 003



Acknowledgment


References

























of






Disease Markers



OncologyJournal of





PPAR Research



Journal of

ObesityJournal of


























of






Disease Markers



OncologyJournal of





PPAR Research



Journal of

ObesityJournal of
















research article mechanism of body weight reducing effect of...

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