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    43 0 A m iC /in Nu i r 199 l;5 3 :430-6 . P rin ted in U SA . 1991 A m erican So cie ty fo r C lin ica l N utrition

    Ene rgy- in ta ke re s tr ic tio n and d ie t-com pos itio ne ffe c ts on ene rgy expend itu re in m en1 2W illia m V R um p ler , J am es L Se a le , C aro lyn W M ile s , a nd C E B o dw e ll3

    ABSTRACT E igh t m en w ere fed a t m ain tenance fo r 2 w k,fo llow ed by 4 w k at 50 % o f m ain tenance , then 1 w k at m ain te -nance . T he d ie ts w ere fo rm ulated to co n ta in e ithe r 40% or 2 0%ofenerg y from fa t. D a ily energ y expend itu re (24-h EE ) w as d e-te rm ined by ind irec t ca lo rim etry a t th e end o fthe 2 -w k m ain te -nance period ; on days 1 , 7 , an d 28 of red uced energ y in tak e;and on days 1 and 7 of re feed ing . D ur in g the reduced-energ ype riod , bo dy w eig h t decreased from 96 .6 to 91 .5 kg and bodyfa t d ecreased from 30 .4% to 27 .7% . T here w ere no s ign ifican td ifferen ces in 2 4-h EE or energy requ irem en ts per u n it bodyw eig h t d ue to d ie t com pos itio n or w eigh t loss. M ain ten an ce m e-tabo l izab le -energy (M E) requ irem en t av erag ed 3 1 .0 kca l/kg bodyw t. O vera ll, the ef ficiency o f M E use rela tive to body s to res w as0 .87 and w as grea te r fo r h igh-fa t th an fo r low -fa t d ie ts. T h erew as som e ev id ence ofan in crease in th e e ffic ien cy o fen ergy u seofb ody sto res a fte r w e igh t lo ss . S u bstra te use ref lected d ie t co rn -pos ition and energy -ba lan ce sta tus indep en den t o f chang es inb od y c om po sitio n. A m J C lin N u tr 1 9 9 l ;5 3 :4 30 -6 .

    KEY W ORDS N utrition al adap ta tion , energy m etab o lism ,body com position , ca lo ric res tr ic tion , d ie t com pos ition

    IntroductionBody fa t am oun ts g rea te r than idea l have bo th neg ative hea lth

    consequences (1 ) and so cia l s tigm as. T he prim ary goa l o f re -stric tio n ofenergy in tak e is th e reduc tion in body fa t m ass . T h isis fo llow ed by the desire to m ain ta in the body a t th is low eredbody fatn es s. C as es o find iv idua ls seem ing ly res istan t to s limm in gor un ab le to m ain ta in w eigh t lo ss a fte r leav in g p resc ribed pro -g rams are num ero us (2 ). Su gges ted exp lan ations fo r the inab ilityo fce rta in ind iv id ua ls to e ithe r lose fat o r m ain ta in fa t loss usua llyinc lu de low m etabo lic ra tes an d g rea te r e ffic iency of use of d ie tenergy . O bese or pos to bese ind iv idu als m ay have low er m etab o licrates than the ir lean coun te rparts (3 -5 ). A low ered m etabo licra te w ould resu lt in low er m ain ten ance energy requ irem en tsand an ap paren t inc rease in th e effic ien cy of en ergy use . T h isw ould con tribu te to an ease ofw e igh t g ain o r re sis tance to slim -m ing . A low ering ofm etabo lic ra te m ay be an ad ap ta tion m ech-an ism to con serve en ergy in respon se to pro lo nged cond itionsof nega tive en ergy ba lance (6 -8 ). T here fo re , in an attem pt toreduce body fa tn ess , ind iv idu a ls w ho restric t en ergy in take m ayincrease the ir res istance to fu rthe r s limm in g and in crease thelik elihoo d o f rega in ing lost w e igh t.

    A no ther fac to r con tribu ting to degree of body fa tness m ay bed ie t com pos itio n . Sev era l p apers (9 -1 1) sugges ted a re lationsh ip

    o f d ie t com pos ition to b ody com po sition in w h ich low erin g fa tleve l in an in d iv idua ls d ie t shou ld resu lt in a low ering of bo dyfa tness . T h is is suppo rted by reports tha t lo ng-te rm m ain tenanceo f in d iv id ua ls on low -fat d ie ts re su lts in a low erin g o f b ody fat-ness (12) .

    T he a im ofth is stud y w as to d ete rm ine the effec t o f reduc tionin energy in take an d the in te rac tio n w ith d ie t com pos itio n o nenerg y expen d itu re , e ffic ien cy of d ieta ry energy use fo r m ain te-nance , su bstra te use , and the com pos itio n of w eigh t loss.

    Sub jects and m ethod sE igh t m ale , m odera te ly overw eig h t su b jec ts w ere se lec ted from

    the genera l pop u lation . T h ey averaged 39 y ofage , 1 8 1 .4 cm inhe igh t, 96 .6 kg in body w eigh t, and 28% body fa t w hen theys ta rted the stud y (T ab le 1) . Each sub jec t co nsum ed th ree mea l sper day pro v ided b y the H um an N u trition R esearch C en te r D ie tS tu dy F ac ility (D SF ) fo r the du ra tion of the s tu dy .

    T he h igh-fat d ie t w as fo rm ula ted to prov id e a com positionrepresen tative of d ie ts norm ally consum ed by the Am ericanpopu la tion (1 3 ) . I t cons is ted o f 14% o f m etab o lizab le ene rg y(M E ) from protein , 40% from fat, and th e ba lance from car-bohydra te . T he low -fat d ie t (1 4% from pro tein , 2 0% o fM E fromfa t) w as fo rm ula ted to redu ce the to tal energy in take from fa tby ha lf. S ub jec ts w ere pe rm itted to consum e on ly the food pro-v ided by th e D SF . T h e ex perim en tal p ro to co l w as app roved bythe H um an S tud ies C omm ittee of the U S D ep ar tm en t o f A gri-cu l tu re -ARS.

    F our su b jects con sum ed the h igh-fa t and fou r, the low -fa t d ie tdu rin g the stud y . T he M E value o feach d iet h ad b een de term in edin a separa te study , o n 42 m en , b y a 7-d to ta l co llec tio n of foo d ,feces , and urine (14). S ix of the su b jec ts in the presen t s tudyhad prev ious ly partic ip ated in lo ng-te rm stud ies in w hich theyw ere fed to m ain ta in body w eig h t. T h e M E in take of th ese pre -v ious stud ies w as used as the m ain tenance lev el o f th is study .

    I From the U S D epartm en t o fA gr icu ltu re , A g ricu ltu ra l R esearch 5cr-v ice , B e ltsv ille H um an N utrition R esearch C en te r, E nergy and P ro te inN u tr ition L abora to ry , B e ltsv ille , M D .

    2 A ddress rep rin t req uests to W V Rum pler, U S D epartm en t o f A g ri-cu ltu re , B e ltsv ille H um an N utrition R esearch C en te r, E nergy and P ro te inN u tr ition L abora to ry , R oom 206 , B uild ing 308 , BA RC -E ast, B eltsv ille ,M D 20705 .

    D e ce as ed .R ece iv ed Ju ly 17 , 1 989 .A ccep ted fo r pub lica tion M ay 10 , 19 90 .

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    10 0

    5 0

    0

    C a lo rim ete r D a y s

    -14 0 1 7 2829 3 5D a y o f E x p e rim e n t

    L A A A A AB od y C om p os itio n A A

    EE A N D EN ERGY -IN TA KE RESTR ICT IO N 431

    FIG 1 . Summ ary of exp er im en ta l p ro toco l, inc lud ing in tak e leve ls and d ay s of energy-exp en d itu re an d body-com -p os it io n d e te rm i na ti on s.

    TA B LE 1Phys ical charac te ristic s o f sub jec ts during in itia l m ain ten an ce periodbefo re w eigh t redu ctionSu bject n um ber A ge Body w eigh t H eigh t Body fa t

    y kg cm %H igh-fa t-d ie t g roup

    I 47 93 178 322 38 1 03 19 6 3 23 52 11 1 18 0 3 84 35 7 3 16 6 27

    Low -fa t-d ie t g ro up1 37 7 7 17 1 3 32 41 10 5 18 7 2 63 35 11 8 18 4 3 74 25 9 3 18 9 1 9

    The 24 -h energ y expend itu re (2 4-h EE ) ofthe tw o sub jec ts w hohad no t partic ipa ted in a prev ious study w as m easured in aB eltsv ille room ca lo rim eter (1 5) befo re the stud y . T he m ain te -nance leve l o fM E in take w as es tim ated as 1 10% ofthe 24 -h EE .T h is estim ate w as based o n prev iou s m easurem en ts (1 5) in w hichEE m easurem en ts w ere m ade on ind iv idu als on lo ng-te rmw eigh t-m ain tenance stud ies . T h e low er EE in the cham ber thanw hen free -liv ing is p resum ed to b e a func tion of th e conf in em entand a red uc tio n in phys ica l ac tiv ity . A ll sub jec ts w ere fed atm ain tenance energy in take fo r 2 w k . T herea fter , energy in takew as red uced to 50% ofm ain tenance fo r 4 w k and then in creasedto m a in tenance fo r 1 w k .

    Tw en ty -four-hour EE w as de term in ed by ind irec t ca lo rim etryby use of the B eltsv ille ro om ca lo rim ete r. S ix m easu rem en tsw ere m ade on each su b jec t (F ig 1) : a t th e end o f the firs t 2 -w km ain tenance period jus t be fo re energ y restric tio n (d ay 0); ondays 1 , 7 , and 28 o f en ergy restric tio n ; and on d ay s 1 (day 29 )and 7 (day 35) afte r re tu rn to m ain tenance energy in take . In

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    ad d ition , tw o sub jec ts w ho had no t been in the ca lo rim ete r fo rat least o ne prev io us 24-h p erio d w ere m easured in th e cham beran ex tra tim e, du rin g th e in itial m a in ten an ce p e riod .

    EE w as ca lcu la ted by th e fo rm ula desc ribed by W eir (16 ) andrates o f use o f subs tra tes (ca rbohydra te , p ro te in , and fa t) w ereca lcu la ted b y use o fth e eq ua tio ns desc ribed by C onso lazio e t a l(17). T hese fo rm ulas u se the da ily p ro duc tion ofca rbon d iox ide ,co nsum ptio n of ox ygen , and excre tio n of u rina ry n itrogen[K je ldah l ( 1 8) A /SN K je l-F oss au tom atic 1 62 10 ana ly zer , F ossE lec tric , H ille rod , D enm ark i. T he fo rm ulas o f W eir (16) andConso laz io e t a l (17) d iffer s ligh tly in the assum ptions ab ou t th eoxygen con sum p tio n and carbo n d iox ide produ c tion assoc ia tedw ith sub stra te ox ida tio n . T h is resu lts in a sm all d iffe rence , < 2% ,in th e ca lcu latio n ofEE d irec tly from oxyg en and ca rb on d iox idev alues o r from substra te -ox ida tion v alues . Fo r purp oses o f sub-s tra te use and ba lan ce ca lcu la tions , d ie t com pos ition (fa t, p ro te in ,and carb ohyd ra te ) w as es tim ated from W att and M errill (19 ) .N o attem pt w as m ade to ad ju st the ox id ation equa tions fo r d if-fe rences b e tw een d ie t and body com pos ition , as sugg es ted byG arlick (20) . T h is w as b ecause of the incom ple te in fo rm ationava ilab le o n am ino ac id , fa tty ac ids , and sug ar com position ofa w ide variety ofcom ponen ts in the d iet. A ltho ugh severa l papers(2 0-23) po in t ou t th a t es tim ates o f subs tra te-use ra tes derivedfrom genera lized equa tions m ay b e sub jec t to apprec iab le e rro rs ,d ep en d ing on the com pos itio n o f the d ie t, severa l w ork ers (9 ,1 1 , 24-26 ) have ob ta ined accep tab le subs tra te-use estim ates byem ploy in g equa tions s im ila r to tho se used in th is stud y .

    S ub jec ts en te red th e ca lo rim ete r a t 0800 and rem ained fo r23 .5 h . In itia l and fina l g as com pos ition s of th e air in the cham berw ere reco rded . D a ta on gas com position an d air f low w ere co l-lected fo r the en tire tim e th e sub ject w as in th e cham ber (1 5 ) .The 23 .5 -h m easured EE w as correc ted fo r d iffe rences in in itia land fina l gas com po sitio n in th e cham ber an d fo r excre tion ofu rina ry n itrogen , and then w as ad jus ted to 24 -h EE .

    W hile in the ca lo rim ete r, each su b jec t fo llow ed the sam e ac-tiv ity p ro toco l (15 ), w h ich inc lud ed 4 h of desk w ork , 1 .5 h of

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    43 2 R UM PL E R E T A L

    * jSEM

    TABLE 2B ody w eigh t, m etab o lizab le en ergy (M E ) in take , 24-h energ y ex pend itu re (24-h EE ), basa l EE , exerc ise EE , resp ira to ry qu o tien t (RQ ), an d urinen itrog en o f m en befo re, during , an d afte r w eigh t loss fed h igh - o r low -fat d ie ts , b y day of experim en t

    D ayO D ay I D ay7 D ay28 D ay29 D ay 35H igh-fa t d iet

    We i g h t ( k g ) 9 7 8 9 7 8 9 4 8 9 2 8 9 1 8 9 1 8ME i nt a k e ( k c a l / d ) 3 0 9 5 9 4 1 5 3 8 4 9 1 5 3 8 4 9 1 5 4 2 4 9 3 0 7 3 9 4 3 0 7 6 9 42 4 - h EE ( k c a l / d ) 2 9 0 2 1 6 0 2 8 7 2 l 5 l . 2 8 6 7 1 47 2 7 5 9 1 3 0 k 2 9 8 8 1 3 9 k 2 9 8 1 5 6 EE, , , k c al / d ) 1 8 8 4 9 5 1 7 7 3 9 5 1 7 0 4 9 4 # { 1 7 6 } 1 6 7 3 9 3 C 1 8 2 6 9 8 1 7 8 2 9 5 EEe z c i . k c a1 ) 2 7 6 6 1 3 0 3 1 4 3 5 5 2 6 3 4 3 3 7 3 2 3 3 4 3 5 0 3 0RQ (C02 /0 2) 0 .86 0 0 .003 0 .835 0 .003 k 0 .80 4 0 .0 06 c 0 .8 22 0 .01 2c 0 .83 7 0 .0 09 k 0 .871 0 .012Ur i ne n i t r o g e n ( g / d ) 1 3 1 1 1 1 1 0 1 1 0 1 1 1 1 1 2 2

    Low -fat d ie tW eig h t(kg) 999 999 979 938 938 948ME i nt a k e ( k c a l / d ) 3 1 8 2 1 0 0 1 5 9 4 5 0 1 5 9 0 5 0 1 5 9 0 5 0 3 1 8 9 5 0 3 1 8 9 5 02 4 - h EE ( k c a l / d ) 2 9 8 9 2 1 8 2 7 9 5 l 6 0 t c 2 7 0 9 l 9 9 2 5 6 1 1 5 1 c x 2 9 3 0 205ab 3 0 3 7 1 9 2 EE, , k c a l / d ) 2 0 4 4 1 1 8 1 8 9 1 9 4 k 1 8 3 9 I 0 8 1 7 9 5 9 7 1 9 0 6 2 b 1 9 7 8 l l 7 EEe , , k c a 1 ) 2 3 8 3 9 2 5 7 2 5 2 7 4 7 1 2 3 9 4 9 3 3 1 4 3 2 7 3 4 1R Q (CO 2O 2) 0 .917 0 .007 0 .881 0 .0 l4 0 .833 0 .009c 0 .840 0 .O lO c 0 .885 0 .01 2 0 .92 5 0 .002Ur i n e n i t r o g e n ( g / d ) 1 6 2 1 6 2 1 1 0 . 5 1 0 0 . 5 1 4 1 1 7 1* . SEM . V alu es w ith in row s w ith sam e su persc rip t le tter do no t d iffe r sign ifican tly , P < 0 .05 .

    m eal consum ption , 1 h of ex erc ise, 7 .5 h of s leep ing , and th eba lance of the tim e spen t as the su b ject desired (eg , w atch ingte lev ision , lis ten ing to the rad io , read ing , desk w o rk). T he 1 hofexerc ise w as d iv ided in to tw o 30 -mm sessions on an exerc iseb icy cle . S leep EE (EE , w as ca lcu lated by m ultip lica tion ofth e ho urly rate o f EE dur in g sleep by 24 h . T he am ount o fenergy ex pended fo r the tw o b icy c ling sessions w asde te rm ined by sub trac ting the s leep ra te from the ene rgy cx -pended during th e exerc ise periods .

    E fficiency ofenergy use (km) and energy requ irem en t (M Ern)as descr ibed by D eB oer e t a l (27 ) w as ca lcu la ted b efo re an d a fte rw eigh t loss as fo llow s:

    R E R E 2k = W , W 2 = effic iency o f u se of d ie ta ry energy _ ef ficiency of use of b ody energy

    W , W 2R E ,

    M EmMEI1 W ,W , W , k m

    w here rne tab o lizab le en ergy in take (M E l) and re ta ined energ y( R E , w hich equ a ls M E l - 24-h EE ) are expressed in kca l/d , andbody w eigh t (W ) is exp ressed in k ilog ram s . S ubscr ip ts 1 an d 2re fe r to m easurem en ts m ade on day 0 and d ay 7 (befo re w eigh tloss) and on day 35 and day 28 (a fter w eigh t loss) . W hen bodysto res and d ie t M E are used equa lly to m eet en ergy requ irem en ts ,km equa ls I .0 . V alues o f km grea te r o r less than 1 .0 rep resen td iet M E being used e ithe r m o re o r less e ffic ien tly , respec tive ly ,than body sto res (2 8).

    B ody w eig h ts o f the sub jec ts w ithou t clo th in g w ere o b tainedonce a w eek afte r a 1 2-h fas t. B ody com pos ition w as de te rm inedby hydrodens ito rn etry [un de rw a ter w e igh ing (UWW ) (29 )] a tth e beg inn ing and end ofthe reduced-energy period .

    S ta tistica l ana ly sis w as perfo rm ed by using th e g en era l-linea r-m ode ls p ro ced ure (G LM ) of th e SA S/STA Tpackage f or p er so na lcom pute rs (SA S Ins titu te Inc , C ary , N C ) . M easurem en t day ,

    d ie t, an d sub jec t n um ber w ere used as c lass variab les. T he vari-ab le day represen ted the m easurem en ts in the calo r im ete r asfo llow s: 0 , day 7 of in itia l m ain tenance period ; 1 , first day o f50% m ain tenance ; 7 , d ay 7 o f 50% m ain tenance ; 2 8 , day 28 o fm ain tenance ; 29 , day 1 o fre tu rn to 100% m ain tenance; and 35 ,

    TABLE 3B od y c om p os it io n chang es an d energy ba lan ce d uring 50%m ain ten ance period on h igh - an d low -fat d ie ts*

    H igh-fa t d ie t(n=4)

    Low -fa t d ie t(n 4)

    Combined(n=8)

    W eigh t (k g)D ay 1 95 .2 16 .4 98 .0 17 .5 96 .6 15 .8D ay 27 89 .9 16 .0 93 .0 17 .0 91 .5 15 .4Di f f e r e n c e - 5 . 2 1 .2 -5 .0 1.2 -5 .1 1. 1

    T o ta l bo dy f a t ( %)D ay I 32 .2 4.3 28 .7 8 .0 30 .4 6. 2D ay 27 29 .6 5 .2 25 .7 8. 4 27 .7 6. 8D iffe rence -2 .5 1 .3 -3 .0 1. 3 -2.8 1. 2

    Fa t-free m ass (kg )D ay 1 6 4. 1 7 .9 69 .6 12 .5 66 .8 10 .1D ay 27 62 .8 8 .6 68 .7 I 1 .9 65 .8 10 .1D if fe rence -1 .3 1 .9 -0 .90 .7 -1 .1 1 . 3

    Tota l bo dy fa t (k g)D ay I 3 1 .1 9 .1 28 .4 10 .8 29 .7 9. 4D ay 27 27 .1 8. 9 24 .3 10 .6 25 .7 9. 2D iffe rence -3 .9 1 .0 -4 .1 1. 0 -4.0 0. 9

    To ta l body energy(Mcal)

    D ay I 353 .1 91 .6 3 33 .5 102 .8 343 .3 90 .7D ay 27 315 .2 88 .6 294 .4 100 .8 304 .8 88 .5D if fe rence -37 .9 7 .6 -39 .1 9.8 -38 .5 8. 1

    T ota l p erc en tbody e n e r g yl o s t a s f a t ( %) 9 6 5 98 2 97 4

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    xx x

    x 0

    0

    EE AND ENERGY -INTAKE RESTRICTION 43 3

    FIG 2. Relationship o f daily energy ex penditure to body w eight inmen before (0 ) and after (X ) w eig ht lo ss .

    TA BLE 4Relative e ffic iency o f ME use for maintenance (km ) and energ yrequirement (MEm) *

    km ME mO ve ral l av erag e (n = 16 ) 0 .87 0.04 31 .0 0. 6High fat (n = 8 ) 0 . 9 3 0 . 0 6 3 1 . 4 1 . 2Low fat (n = 8 ) 0 .82 0 .04 t 30 .6 0 .6 tB e fore w e ight lo s s (n = 8 ) 0 .94 0 .05 30 .3 0. 8A fter w eig ht lo ss (n = 8) 0 .81 0 .0 S f 32 .5 0 .9 H ig h-fat-die t g ro up

    Before (n = 4 ) 0 .99 0.0 8 30 .3 1. 7After (n = 4 ) 0 .87 0.0 5 32 .6 1 .6

    Low -fat-die t groupBefore (n = 4) 0 .90 0.0 6t 2 9 .9 0 .4After(n=4) 0 .75 0 .0 6 t 31 .21 .6* i5EMt S ignificantly different from corresponding high-fat-die t gro up, P

    . 2 8 0 0(2wz 2 6 0 0 -

    6 0 7 0 8 0 9 0W E IG H T (k g)

    1 0 0 1 1 0 1 2 0

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    43 4 R UM PL E R E T A L(T ab le 5 ). H ow ev er , t h e r e w as n o sign if ican t in t er ac t ion b e tw eend ie t com position and w eigh t loss on subs trate use .

    B ot h t h e h igh - and low -fa t g rou ps w ere in pos itiv e pro tein ,ca rbohydra te, and fa t b alance w hile in th e ca lo rim ete r con sum ingthe w eigh t-m ain tenance in take lev el befo re and afte r w eigh t loss .H ow ev er , car b oh y d r a t e b a lan ce w as sign if ica n t ly low er in t h eh igh -fa t g rou p a fte r w e ig h t lo ss .

    Afte r 7 d at 50% ME l, p ro te in ba lance w as near ze ro an d fa tba lance w as m arked ly nega tive in bo th d ie t g roups bu t w as no tsign if ican tly d iffe ren t b etw een d ie t g roups . C arbohydra te balancew as n ega t iv e f o r t h e h igh -f a t g r ou p a n d p osit iv e f o r t h e low -f a tg r o u p .

    Som e sign ifican t d iffe ren ces in use and ba lan ce occurred onthe firs t day ofreduced in tak e (day 1) an d the firs t day of re tu rnto m ain tenance afte r w eigh t loss (day 29) w hen com pared w ithm easurem en ts m ade at the sam e in take leve l. O n day 1 w hencom pared w ith day 7 or 28 , carbohyd ra te u se w as g reate r andba lance w as m uch m ore nega tiv e; fa t u se w as low er and b a lancew as m uch less nega tive ; p ro te in ba lance w as m ore nega tive onthe low -fa t d ie t th an on e ithe r the h igh -fa t d ie t o r d uring anyo ther m easurem en t m ad e du rin g the s tudy . O n day 29 w hencom pared w ith days 0 and 35 , p ro te in b alance w as h igh er, fa tba lance w as low er, and carbo hydra te ba lan ce w as low er. In ad-d itio n , ca rbo hydra te ba lan ce w as sign ifican tly m ore pos itive onthe low -fa t d ie t th an on the h igh-fa t d ie t a fte r w eigh t loss.

    D iscuss ionE E d ec lined d uring the reduced-en erg y-in take period . H ow -

    ever, M E req u irem en t w as n o t a ffec ted b y w eigh t loss . In s tud iesw ith w om en , D eBo er e t a l (27) repor ted a low er EE and a low er

    T A B L E 5

    M E r eq u ir em en t a f t e r w eig h t lo s s b u t t h e d if f e r en ce w as n o tsig n ifican t. R avu ssin e t a l (30 ) a lso fou nd no red uc tion in E Ew ith w eigh t loss tha t cou ld no t be exp la ined by red uc tion s inbody m ass , energ y in take , o r low ered ac tiv ity . S om e stud ies haved em onstra ted or im plied a d iffe rence in energy requ irem en t a fte rw eig h t loss. G eissle r e t a l (3 ) found a 1 5% low er EE in pos tobesethan in lean ind iv idua ls m atched by body com po sition . H ow ever,m easurem en ts w ere m ade on the pos tobese ind iv idua ls w hilethey w ere in nega tive energy ba lance, w hich w ou ld tend to low erthe m etabo lic ra te . A lso , it can no t be de term in ed ifthe po stobeseind iv idua ls had a low er m etabo lic ra te befo re w eig h t lo ss . S evera ls tu d ies rep ort a reduc tion in res ting EE in respo nse to rap idw eig h t loss (6 -8 ). T hese stu d ies use very -low -ca lo rie d ie ts ( 800kca l/d ) to ach ieve w e igh t lo ss . Th ese ve ry low en e rgy in tak esresu lt in sign if ican t loss o f FFM and m ay no t be com parab le tositu atio ns on slow w eigh t lo ss w ith m in im a l ch anges in FFM .

    D iet d id no t a ffec t 24 -h EE or en ergy req u irem en ts. H ow ever,sleep EE ten ded to be grea te r on th e low -fa t than o n the h igh -fa t d ie t (T ab le 2). A probab le exp lana tion fo r th is is th a t th einc rease in E E assoc ia ted w ith the m ea l (the rm ic effec t) w aspro lon ged on the low -fat d ie t, pe rhaps lasting u n til th e nex tm orn ing . T h is w ould e leva te EE du rin g sleep , w hich w as th ebasis o fthe estim ate o f s leep EE , bu t it w ou ld no t a ffec t the 24-h to ta l. H urn i e t al (31) a lso repo rted h igher EE during s leepbu t no t o ver 24 h on a low -fa t d iet vs a h igh -fa t d ie t.

    T h e effic iency of d iet M E use re la tive to body s to res w a sm arked ly d ifferen t be tw een d iets an d af te r w eig h t loss. T h ish igher-e ff ic iency v a lu e ofthe h igh-fa t d ie t im plies tha t a t o r be lowm ain ten an ce, h igh-fa t d ie ts w ill tend to be m ore ef fic ien tly usedfo r m ee ting m ain tenance req u irem en ts th an low -fa t d ie ts , re -qu irin g s lig h tly less energy in take of th e h igh-fa t d ie t fo r w eigh t

    T o ta l su bstra te use and n et subs trate ba lan ce befo re , during , and af te r w eigh t loss in m en on h igh- and low -fat d ie ts, by day of experim en tD ayO D ayl D ay7 D ay28 D ay29 D ay35

    g/ dPro te in

    U seHi g h - f a t d i e t 8 4 7 6 0 8 6 1 8 6 1 8 7 0 6 7 3 1 2 L ow- f a t d i e t 1 0 0 1 2 t 8 9 1 6 7 2 c t 6 4 2 c t 9 0 5 t l 0 6 6 t

    Ne tHi g h - f a t d i e t 3 1 6 2 l O - 3 4 . 1 4 5 6 4 2 4 L ow- f a t d i e t 2 8 14sb 2 5 2 d t 3 2 c 1 2 . 7 c 3 9 5 k 2 2 6 t

    Carbohydra teU se

    Hi g h - f a t d i e t 3 4 7 2 5 a b 2 9 0 1 5 2 1 6 1 9 2 5 1 3 2 & 3 ( ) 3 o C 3 8 6 3 3 L o w- f a t d i e t 4 8 8 3 9 t 3 7 9 4 4 t 2 6 4 2 7 c 2 6 6 2 5 c t 4 0 3 3 l t 5 l 4 33 t

    Ne tHi g h- f a t d i e t 2g 2 5 a b - 1 0 0 14 d - 2 6 l 8C - 6 2 3 1. l 6 8 3 1 - l 2 3 3 L o w- f a t d i e t 7 5 2 7 - 9 8 3 8 1 8 2 3 C 1 5 1 9 . 8 c t 1 5 9 2 0 t 4 9 1 8

    Fa tUs e

    Hi g h - f a t d i e t l l s 3 c 1 4 8 11 b 1 7 9 1 3 1 5 2 10 b l 4 8 I Ob l l l 1 3 cL o w- f a t d i e t 5 5 l 3 Ct 8 8 l s b t 1 3 9 1 7 t 1 2 4 l 4 t 9 1 l 8 t 4 6 6 Ct

    Ne tHi g h - f a t d i e t 3 0 5 3 4 1 4 L o w- f a t d i e t 1 2 1 4 - 5 4 l 4 C - 1 0 5 1 7 - 9 0 1 4 - 2 4 l 9 2 1 6

    S S E M . V alues w ith in row s w ith d iffe ren t superscr ip ts d iffer sign ifican tly , P < 0 .05 .t Sign if ican tly d if fe ren t from high-fa t-d ie t va lu e, P < 0 .05 .

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    EE A ND ENERGY -IN TAK E RESTR ICT IO N 43 5m ain ten an ce . V an E s e t a l (28 ) rep orted th e effic iency va lues o f0 .8 7-0 .94 on a d ie t s im ila r in com position to the h igh-fa t d ie to f th is s tudy . T he low er e ffic iency , o f M E for m ain tenance re l-a tive to body s to res, a fte r w eigh t lo ss sugg es ts tha t ind iv id ua lshav ing los t w e ig h t use bo dy sto res m ore eff ic ien tly than they d ob efo re w eigh t loss . D eBoer e t a l (27) repor ted ef ficiency va luesm easu red in w om en tha t w ere h igher a fte r w eig h t loss than b efo rew eigh t lo ss and w ere low er th an the effic iency fo r m ain tenanceva lues o f the h igh-fa t d ie t in th is s tudy .

    A s po in ted o u t ea rlie r, the es tim ates o f sub stra te ox ida tionan d ba lance m ust b e v iew ed w ith an aw aren ess o f the ir lim ita -tion s. H ow ever, th e genera l pa tte rn and even th e m agn itude ofthe chang es in su bstra te ba lance are re flec ted in th e changes inbody com position . D urin g th e w eigh t-loss phase of the stud y ,by da y 7, pro te in an d carboh ydra te b a lance w ere no t sign ifican tlyd if fe ren t from zero w hereas fa t ba lance w as m arked ly n eg ative .H ow ever, FFM in th e h igh -fat g roup ten ded to decrease m orethan in th e low -fa t g roup , w hich is re flected in the m ore nega tivecarb ohydrate ba lan ce in the h igh -fa t g roup . S im ila rly , lo sses o ffa t m ass ten ded to be g rea te r in the low -fa t g ro up an d fa t subs tra teb a lan ce w as m ore nega tive .

    Subs tra te use and ba lance genera lly ref lec ted en ergy ba lance .W hen sub jec ts w ere fed at a w eig h t-m ain tenan ce in tak e , be fo rew eigh t loss , subs tra te use re flec ted the d iet com position , w iththe h ig h-fa t g roup u sing m ore fa t and less ca rboh ydra te da ilythan the low -fa t g ro up . A f te r the sub jec ts had been fed a t anen ergy in take leve l o f 50% of m ain tenance fo r 28 d , subs tra teuse s till re flec ted d ie t com pos ition . H ow ever, fa t use inc reasedan d carb ohydrate use decreased in bo th groups , re flec tin g th eincreased con tribu tion of body fa t sto res in m ee ting the m d i-v idu als energ y requ irem en ts.

    U pon re feed ing (re tu rn to 100% m ain tenance leve l a fterweigh t- loss period ) fo r 7 d (day 35), p ro te in , ca rb ohydrate , andfa t ba lance w ere no t s ign ifican tly d iffe ren t from befo re the w eig h t-lo ss p e r iod . H ow ev er , ca r b o h y d r a t e b a lan ce t en d ed to b e low eran d fa t ba lance tended to be h igher a fte r w eigh t lo ss than befo rew eigh t loss, sug gesting a fat-sparing effec t o f w eigh t loss.

    O v erall, su bstra te use re flec ted d ie t com pos itio n a t in takesb o th near m ain tenance an d a t 50% of m ain tenance . T he changein bo dy com positio n co rre spon d ing to the lo ss o f bo dy weigh th ad n o in fluence o n sub stra te use ou ts id e o fthe con tribu tion o fbody fa t to the en ergy-subs trate poo l. In a series o f p ap ers F la ttan d cow orkers (9 - 1 1 ) sug gested th a t chang es in in tak e o f ca r-b ohyd ra te and fa t in fluence th e o x ida tio n rates o fthese subs tra teso n ly rela tive to th eir body sto res . Sm all changes in carboh ydra tein take resu lt in corresp ond ing inverse ch an ges in carb ohydrateo x ida tio n ra tes becau se of the lim ited capac ity o f the body tos to re ca rboh ydra te . H ow ever, ox id a tion ra tes o ffa t a re re la tive lyinsens itive to changes in in tak e o f fa t because of the la rg e s to reso fenergy ava ilab le from fa t. T h is im plies short-te rm energ y def-ic its w ou ld be com pensa ted fo r by dep le tio n of ca rb ohydrates to res , and fa t m ob iliza tio n w ould be slow er to respond . In th iss tudy , w hen 24-h ox id ation ra tes a re com pared be tw een days 0(las t day of m ain ten an ce) and I (firs t day of reduced ca lo ries),fat ox ida tion ra tes h ad increased m arked ly an d carb ohydrateox ida tion had dec lin ed nearly the sam e am ount (on an energybasis rathe r than on a m ass bas is ). T he opposing respo nse is seenw hen d ay 28 (las t day of reduced energy in take) an d d ay 29(first day of m ain tenance afte r w eigh t loss ) are com pared . Fa tox ida tion dec lines and carb ohyd ra te ox ida tio n in creases in re -spo nse to grea te r d ie ta ry en ergy in tak e. T here is som e support

    fo r th e s low er respon se of body fa t sto res to chan ges in en ergyin take . C arbohydra te ox ida tion w as grea te r on the first day ofred uced energ y in take (day 1 ) and s to rage w as grea te r on thefirs t day of the re feed in g a t m ain tenance (day 29) than duringsubseq uen t d ay s at the sam e M E in take .

    In th is experim en t, a s ign ifican t redu c tion in EE occu rredaf te r the redu ction of M E t and during the w eigh t-loss period .H ow ever, energy requ irem en ts on a b ody -w e igh t basis w ere n o tsign ifican tly e ffec ted b y w eig h t reduc tion . T h is does no t re fu teobserv a tion s of low er m etabo lic ra te in obese o r po stobese in -d iv id ua ls (3 , 4 ). H ow eve r , it do es sugg es t tha t ifo bese ind iv idu alsh av e low er m e tabo lic ra te s , it p ro bab ly is no t a func tion of recen tm oderate w e igh t lo s s .

    T he re w ere sign ifican t d ifferences in e ffic iency o f use o f d ie tM E r e la t iv e t o body sto res . T he h ig h -f a t d ie t w as m or e ef f ic ien t lyused than w as th e low -fa t d iet re la tive to b ody sto re s. T he e ff i-ciency of M E use re lative to body s to res w as low er a fter w e igh tlo ss in bo th d ie ts , sug gesting th at e ithe r bod y sto re s w ere m oree ff icien tly used a fter w e igh t lo ss th an b efo re o r th at the e ff icien cyof use o f M E dec reased . If bod y sto re s w ere m ore e ff ic ien tlyu sed , th is w ou ld con tr ib u te to a res istance to s limm in g w h ile inn eg ative en ergy ba lance w ith ou t ch ang ing th e energy requ ire-m en ts o f the ind iv idu a l.

    In th is s tudy , th e ra te o f fa t o x ida tion re spo nded w ith in 1 dto ch an ges in ene rg y sta tus. T he re w as no ev id en ce of largechanges in carboh ydra te s to res in response to sh ort-term (l-d )changes in energy ava ilab ility . C lear ly , in th is stud y , the com -positio n of su bstrate used fo r m ee ting ene rg y needs ref lects en -ergy-b a lance s ta tus and d iet com pos ition and w as n o t m arked lyaffec ted by changes (w eig h t lo ss ) in body s to res. 0R e f e r e n c e s1 . D epartm en t o f H ea lth and H um an Serv ices. T he Surgeon G enera ls

    repo rt on nu tr ition an d hea lth . W ash in g ton , D C : US Governm en tPrin ting O ffice , 19 88 . [D H H S publication (PH S ) 88 -50210 .)

    2. M iller D S , P arso nage S . R es istance to slimm in g : ad ap tation or il-l u si o n? La nc et 1 97 5; l : 7 7 3- 5 .

    3 . G eissle r CA , M ille r D S , S hah M . Th e da ily m etabo lic ra te o f thepos t-obese and the lean . Am J C lin N utr l9 87 ;45 :9 l4 -20 .

    4 . R avussin E , L illio ja S , K now ler W C , e t a l. R educed ra te o f energyexpend itu re as a risk fac to r fo r body w eigh t ga in . N EngI J M ed1 9 88 ; 3 l 8 : 4 6 7 - 7 2 .

    5 . Sh ah M , M iller D S , G eiss le r CA . L ow er m etabo lic ra tes o f po stobese versus lean w om en: the rm ogenes is, basa l m etabo lic ra te andgene tics . E ur J C lin N utr l988 ;42 :7 4 l-52 .

    6 . H ill JO , Sp arling PB , S he ild s TW , H elle r PA . E ffec ts o fexerc ise andfood restric tio n on body com po sitio n and m etabo lic ra te in obesew om en . Am J C lin N u tr 198 7 ;46 :622-30 .

    7. W eigh le D S , Sande K J , Ive rius PH , M onsen ER , B runze ll J D . Weigh tlo ss leads to a m arked d ecrease in no nres ting ene rgy expen d itu re inam bula to ry h um an su b jec ts . M etabo lism l9 88 ;37 :930 -6 .

    8 . E llio t D L , G o ld berg L , K ueh l K S , B enne tt W M. Sus tained depressionof the res ting m etabo lic ra te af te r m ass ive w eigh t loss. Am J C linN u tr 1 98 9; 49 :9 3- 6.

    9 . F la tt JP , R avussin E , A ch eson K J, Jequ ier E . E ffec ts o f d ieta ry fa ton postp ran d ia l subs tra te ox ida tion and on carboh ydra te an d fa tba la nc es. J C lin In ve st l98 5;76 :lO l9 -2 4.

    10 . F la tt JP . T he d ifferen ce in the s to rage capac ities fo r carbohyd ra teand fo r fa t and its im plica tion s in the regu la tion o f body weigh t .A nn N Y A cad Sc i l 9 8 7 ; 4 9 9: l 0 4 - 2 3 .

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