resistant starch - the effect on postprandial glycemia, hormonal response, and satiety

8/13/2019 Resistant Starch - The Effect on Postprandial Glycemia, Hormonal Response, And Satiety

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54 4 Am J C /in N u ir 19 94 ;60 :54 4-5 I . P rin te d in U S A . 1994 A m eric an S o cie ty fo r C l in ica l N utritio n

R es is tan t s ta rch : the e ffe c t on p os tp rand ia l g lycem iaho rm ona l re sponse and sa tie ty13A nne R ahen , An na T ag liab ue , N ie ls J C hris ten sen , Joo p M ad sen , Je tis J H o is t, and A n te A s trup

A B S T R A C T T h e effect o f resistan t s tarch (R S ) o n po stp ran -d ial p la sm a co nc en trations o f g lu co se, lip ids , and ho rm o ne s, a ndo n sub jec tiv e sa tie ty and pa la tab ility ra tings w as in vestiga ted in10 hea lthy , norm al-w eigh t, yo ung m ales . T he tes t m ea ls con-

sisted o f 50 g p reg ela tin ized sta rch (0 % R S ) (5 ) on 5 0 g rawpo ta to s ta rch (54% R S) (R ) tog ether w ith 5 00 g art ific ia lly sw e et-ened syrup . A fte r the R m eal p ostp ran d ia l p lasm a concen tra tionso f g luco se , lac ta te , in su lin , gas tric in h ib ito ry po ly pep tide (G IP),g lu cag on-like pep tide - I and ep ineph n ine w ere s ig n ifican tlylow er co m pared w ith a fte r the S m ea l. M o reover, sub jec tiv esco res fo r sa tiety and fu llness w ere sign ifican tly low er a fte r theR m eal than after th e S m ea l. D iffe rences in G IP , tex tu re , andpa la tab ility m ay have b een in vo lved in these f in d ings . In conc lu -sion , th e rep lace m en t of d iges tib le s tarch w ith R S re su l ted insign ifica n t re du ctio ns in p ostp ran d ial g lycem ia an d in su linem ia ,and in th e sub jec tiv e sensa tions of sa tie ty . A m J C liii N u trI 994;60:544-5 I.

K Y W OR S P otato sta rch , app etite , p ala tab ility , lac ta te ,insu lin , ga stric inh ib ito ry p oly pep tid e, catech o la m ines

Introduc t ion

U ntil recen tly , stanch w as be liev ed to be 1 00% digested in thesm all in tes tin e- independ en t o f the source , typ e, and p repara tio nof the s tarch . H o w ever, w ith in the pas t 10 y it has been foun dtha t, desp ite the fac t th at pancrea tic a -am y lase is p resen t in thegu t in am ple am ou n ts ( I ), a fra ctio n of the ing este d sta rch pas sesund ig es ted to the la rge bow el (2 -4 ). H ere the s ta rch underg oesa m o re o r less co m plete fe rm en ta tion , resu lting in the produc tionand u p take of short-cha in fa tty acids (ace tic ac id , b u tyn ic ac id ,and prop io n ic ac id ) (5 , 6 ). T h is frac tion of the s tarch has beennam ed resis tan t sta rch (R S) and h as b een d ef in ed by the E uro-pean F L A IR C on certed A ction o n R esistan t S ta rch (E U R E ST A )as the su m of s ta rch an d produ cts o f s ta rch hyd ro lys is n o t ab -so rbed in the sm all in tes tine o f hea lth y ind iv id ua ls .

T h e am o unt o f R S p resen t in s ta rch - rich foo ds depend s onsevera l fac to rs, ic , the sou rce, r ip en ess , p rocess in g , p repara tion ,and sto rag e of the fo ods . It has been sh ow n tha t stanch from w hiteb read , porridg e o a ts , and corn f lakes is a lm os t co m ple te ly d i-gested in the sm all in testine (4 ) w h ereas na tive sta rch fro m ba-nana and un co oked po ta to is h igh ly resis tan t to hy dro lys is in v itro(7 ) an d in v ivo (8). W he n th e p otato is c oo ked the starc h granu lesge la tin ize and becom e read ily d ig es tib le, w hereas coo ling o f thepo ta to reverses th is ge la tin iza tion process and ren ders I 2% ofth e sta rch res istan t to sm all in tes tina l d ig es tio n (9 ).

T he im p ortance of R S to hum an hea lth , ie , d iab e tes , ov er -w eig h t, ca rd iov ascu la r d iseases , o r cancer, is s till n o t know n.H ow ev er, because R S -un like nonres istan t stanch-is n o t d i-ge ste d an d th ere fore n o t a bso rbe d as g luc ose in th e sm a ll in tes tin eo f h ealthy h um ans (8 , 9 ) , a reduc tion in bo th pos tp rand ia l g ly -c em ia an d in su l ine m ia c an b e exp ec ted afte r th e in ta ke o f R Scom p ared w ith d iges tib le s ta rch . Such an effec t o f R S m ay bebe ne fic ia l in the con tro l o f d ia betes. M ore ove r, R S m ay th rou ghm ech anism s sim ila r to tho se e xer ted by so lub le d ieta ry f iber in -fluence the am ou n t and rate o f absorp tion o f o ther nu trien ts inthe d ie t, ie , g lucose and fa t, w h ich m ay be benefic ia l in th e con -tro l o f g ly cem ia or lip idem ia.

T he po ten tia l use of R S as a w eigh t-redu c in g agen t m ay a lsobe of in teres t becau se th e energ y v alue of 1 g R S , in clud in g thecon tribu tion from ferm en ta tion p roduc ts , has b een es tim ated tobe on ly 9 .0 -9 .8 kJ /g , ie , ha lfthe va lu e ofd ig es tib le s tarch (10).It is th ere fore tem pting to su gg est R S a s a w e igh t-lo ss ag en t;ho w ever, the im p ac t o f R S o n m acno nu trien t ba lance and appe titecon tro l m ay be less benefic ia l to w eigh t regu lation . T h is hyp o th -es is is ba sed on th e rece n tly d eve lop ed co nce p ts o n th e regu lationof energy and m acnon u trien t ba lance. T hus , the re seem s to b e ac lose regu la tion be tw een the bod ys m acronu trien t s to res and theth ree m acronu tn en ts (fat, p ro te in , an d carbohy dra te) in the d ie t(1 1 , 12) . T h e m ain tenance of the bo dys re la tiv ely sm all ca rbo -hy dra te ston es seem s esp ecia lly c ru cia l to overa ll energy b alance I I . M oreover, recen t s tud ies h av e show n tha t the co n tro l o fappe tite m ay b e in fluenced by the am ount o f av a ilab le ca rbohy -d ra te in th e d ie t ( 1 3, 1 4) and tha t con sum p tion of a cer ta in am oun to f ca rboh ydra te (o r p ro te in ) in a m ea l m ay be necessa ry toach iev e satia tio n (15 ) . Inc reasing the am ou nt o f R S in th e d ie tw ith a re su ltan t d ecreased am ou nt of ab so rbe d carboh yd rate a nda dec rea sed c arb ohy dra te- to - fa t ra tio in th e d ie t m ay thu s lead to

Fro m th e R esearch D epartm en t o f H u m an N utrition . C en tre fo rF o o d R esearch . T he R oya l V ete rina ry and A gricu ltu ra l U nivers ity .F re de niksbe rg . D e nm a rk: the D epa rtm e nt o f H um an N utrition . U niv er-sity of P av ia, Italy : the D ep artm e nt of In te rn al M edic ine and E ndoc rin-olo gy . H en ley H o spita l. D enm ark : a nd the D e pa rtm ent of P h ysio log y .T he P an um Institu te , U nivers ity of C o pen ha gen . D e nm ark .

2 S uppo rted by gra nts from T he D anish R esea rc h a nd D ev elop m en tProg ram m e for F ood T ech nology 19 90 - 1994 and the D an ish M edica lR esearch C oun cil g ran t I 2 -9537 -3 .

A dd res s re prin t req ues ts to A R a ben . R ese arch D ep artm e nt ofH um an N u tr ition , C en tre fo r F oo d R esearch , T he R oya l V ete rina ry andA gric u ltura l U niv ers ity . 25 R olighed sv ej. D K -1 958 F rc deriksbe rg C .

R e ceived N ove m b er 2 . 1 993 .A ccep ted fo r p ub lica tion M ay 9 . 19 94 .


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R E SIS T A N T S T A R C H , G L Y C E M IA . A N D SA T IE T Y 54 5T A B L E IS ub je ct c ha ra ct er is ti cs

D e n m a r kii = 5)

I ta ly = 5)

Al l = 10 )

A ge (y) 25 .6 1 .9 21 .2 0 .7 23 .4 1 .2H e ight (m ) 1 .80 t 0 .01 1 .7 8 0 .02 1.79 0 .01W eig ht (k g) 71 .2 3.0 74.0 3 .5 72.6 2.2B M I2 22 .0 1 .1 23.2 0 .6 22.6 0.6F a t ma s s ( 1 8 . 6 2 .4 20 .8 1 .3 19 .7 1 .3

. S E M . In k g/m .

a red uct ion in th e sat ia ting p ow er of the d ie t. T h is m ay resu lt inov enconsum ption an d subseq uen t w eigh t g ain . H ow ever, if R Sh as a dietary-fiber-like po sit ive ef fec t o n satie ty ( I 6 - 1 8) anin cre ase in the am ou nt of R S in the d iet m ay be ben efic ia l tow eig h t m ain te nan ce.

T he purp ose of the p resen t stu dy w as to inves tig ate th e acu tee ffec ts o f R S vs d ig es tib le non resistan t sta rch on pos tp rand ia lc han ges in g ly cem ia , ho rm ona l re spo nse, an d su bjec tiv e se nsa -tion s o f hu nge r an d sa tie ty . T he tw o test starche s w ere g iv en inas pure a fo rm as possib le to c la rify the ro le o f the sta rches p ense w itho u t the in terfe ren ce f rom oth er nu trie n ts .

Sub jects an d m ethodsT he experim en t w as perfo rm ed as a jo in t stud y w ith in the

E U R E S T A fram ew ork . E xac tly th e sam e experim en t w as con-du c ted a t tw o cen te rs: the R esearch D epartm en t o f H um anN utritio n , T h e R oy al V ete rina ry and A gricu ltu ra l U nivers ity ,D en m ark , and th e D epartm en t o f H um an N utrition , U nivers ityo f Pav ia , Ita ly .

ubjectsIn each cen te r five hea lthy m ale sub jec ts (20-31 y of age ,

no rm al-w eigh t, nonsm okers , n o t e lite a th le tes ) w ith no h is to ry ofo bes ity or d ia betes, p art ic ip ate d in the stu dy (T ab le 1 ). F em alesw ere no t inc lu ded to avo id possib le d iffe rences due to th e m en-s trua l c ycle . T he su b je cts ene rgy n eed s du rin g the stud y w erede te rm ined b y us in g W H O tab les accord in g to age , w e ig h t,he igh t, an d sex . T he m ultip lica tio n fac to r 1 .78 w as used to ac-cou n t fo r m ed ium phy sica l activ ity lev el o f th e su b jec ts (19). T hebo dy com p osition of the sub jects w as estim ated by the b io im -pedance m etho d by an A nim ater (H T S E n gineerin g Inc . O dense)in D en m ark and B IA 109 (R il S ys tem s , D etro it) in Ita ly . Fa t-freem ass w as c alcu lated b y u sin g the eq uat ion o f D eu ne nbe rg et a l(2 0). T h e s tud y w as app rov ed b y the M un icipal E th ica l C om -m itte e o f C op enh age n and F red en iksb eng to be in a cco rda nce w iththe H elsink i-Il d ec la ra tio n and all su b jects gav e w ritten con sen ta fte r the experim en ta l p ro ced ure had been ex p la in ed to them .

ietsT h e tw o tes t m ea ls con sisted o f 50 g raw po ta to stanch (54 .1%

R S ) (R ) on 5 0 g prege la tin ized po ta to sta rch (100 % d ig es tib le)(5 ) m ixed in to 500 m L dilu ted artific ia lly sw ee tened fru it sy rup a b le 2 ). T he sy ru p w a s b ase d on ap ple , g ra pe. re d c urren t,e lde rbe rry , b la ck cu rre n t, an d che rry a nd com m erc ially pun -c hased fro m I r m a , D en m ark . T he s ta rches w ere produced and

T A B L E 2E U R E ST A refe rence sta rch m ate ria ls

Fo od D M

C al c u la ted s tarc h v alue s

T S R D S S D S R S% (;y z

R aw p otato s tarch 83 .4 81 .3 6 .0 21 .2 54 .1P r e g e l a t i n i z e d p o t a t o s t a r c h 9 5 . 1 9 2 . 9 8 7 . 9 5 . 2 - D M , dry m atte r: T S . to ta l s tarc h : R D S . rap id ly d ige stib le sta rch :

S D S , slow ly d iges tib le sta rch : R S . resis tan t sta rch . A na lyzed from th eD unn C lin ica l N utritio n C en tre . M ethod s descr ib ed by E n g lyst e t al (21)and E ng lyst and C um m ings (22 ).

su pp lied from lIns titu t N ationa l de Ia R echerche A gron om ique(IN R A ), N an tes, F rance . B o th sta rches appeared as a w hite flou r.iu s t be fo re co nsu m ption th e s ta rch w as s low ly p oured in to thesy rup w h ile m ix ing it w ith a h an d m ix er at th e slo w e st spe ed toav oid an inc rea se in tem pe ratu re o f th e m ix tu re . S ev era l o therw ay s o f ad m in is tra tion w ere tes ted befo re d ec id ing on th ese testm ea ls . H ow ever, the p rege la tin ized stanch cou ld no t be m ixedin to o ther liqu id s w ithou t p roduc ing a v ery un p leasan t, ined ib lesu bstance . B ecause h ea ting w o uld affec t the ph ysica l p ropertieso f th e raw starch , a co ld -m ea l p repara tion had to be used . T hesta rches co u ld thus no t be baked in to a b read or used in a pas ta .T he best com pro m ise fo r these tw o sta rches w as th erefo re to m ixthem w ith the sy rup . F or the R m eal th is resu lted in a d rinkab lem ix tu re w hereas the S m eal w as a porrid ge-lik e ge l to be ea tenby spoo n . A lth oug h the sy rup w as sw eetene d w ith a rtific ia lsw ee ten ers it con ta ined som e fru i t su gars natura lly o ccu rring inthe berries and fru its (T ab le 3 ).

E ach tes t d ay w as p receded by 3 d on an iden tica l canb ohy-d ra te -n ich d ie t (60 % of energ y as carbo hydra te , 28% as fa t, an d12 as pro tein . and 3 .5 g d ieta ry fib er/M i) , p rep are d at th e d e-

pa rtm en ts fro m foo d item s accord ing to each sub jec ts in d iv idua lenergy requ irem en ts . and ad justed to the n eares t 0 .5 M i. T hesu b jects w ere ins truc ted to ad here stric tly to th e d ie t. If sub jec ts

T A B L E 3C arbo hydrate an d energy c onten ts in th e tw o tes t m ea ls(50 g pota to s ta rc h + 50 0 m L d ilu te d f ru it s yru p)

T es t m ea lR S

T o tal sta rch in 5 0 g po tato starch (g) 40 .7 46 .5R S (g) 27 .1 0 .0R D S (g ) 3.0 4 4 . 0SD S(g) 10 .6 2 .6F ruit suga rs fro m syrup (g) 8 .4 8 .4

G lucose (g ) 2 .9 2 .9Fru c to se (g ) 4 .4 4 .4S u cro se(g ) 1 .1 1 .1

T ota l d igestib le ca rboh ydra te (g )2 22 .0 5 4 .9T o ta l energy (k l)3 367 9 17

R , raw pota to s ta rch : S . p rege la tin ized po ta to s ta rch : R S , resis tan ts tarch : R D S . rea dily dige st ib le sta rch : S D S , slo w ly dig es tib le s tarc h .

2 N o t tak in g fe rm en tation in to conside ra tion .3 1 6. 7 k J / g .


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54 6 R A B E N E T A Lcou ld no t consum e a ll th e h od , they had to bring the le ftoversto the d epartm en ts fo r w eigh ing and reg is tration . T he sam e foodw as d ed uc ted from the d ie t during the fo llow ing preexpen im en ta lpe riod s. T he sub jects w ere ins truc ted to ab sta in f rom stren uousph ysica l ac tiv ity fo r the 2 d befo re the test d ays. T og ether w iththe s tand ard d ie t th is sho u ld ensure equa lly f illed g ly cogen s to nesand s im ila r m acnonu tn ien t b a lance on the 2 test days . T he com -pu te r d a tabases o f foods f rom the N ationa l F ood A gency ofD en m ark (D an kos t) and Italy (IN N ) w ere u sed in the ca lcu la tio nso f energ y and nu trien t com pos itio n o f the test d ie ts.

E .qeriitie iz ta l p rok )co /T he tw o tes t m ea ls w ere g iven in a crossov er design on se-

para te days w ith I w k and no m ore than 6 w k separa ting thetest days . O n the test day the su b jec ts a rrived a t the ins titu te at0 80 0 , w ith a m in im um o f ph ys ical act iv i ty , by ca r, bu s, o r tra inafter ha vin g fa sted for 1 2 h from the e ven ing b efo re . A fter sub -jects vo id ed and w ere w eighed (to the nearest 1 00 g), b io im ped-ance w as m easu red . T h e sub jec ts th en res ted in the sup in e p o-s ition on a bed covered w ith an an tidecub itus m attress w ith sligh te leva tion of the head . A V enflon ca th e te r (V igg o . G o th enborg .S w eden ) w as inse rted in an an tecub ita l a rm ve in . A fter a 10-m mres t a fas ting b lo od sam ple w as tak en and afte r a fu rthe r 20-m mres t nesting m etab o lic ra te w as m easu red by ind irec t ca lo rim etryby us ing a v en tila ted hood . A second fasting b loo d sam ple w astaken h erea fte r an d the tes t m eal w as then served an d con sum edw ith in 1 0 m m . E xac tly the sa m e tim e w as spen t on th e tw o testm eals fo r each ind iv idua l sub jec t. P os tp rand ia l en ergy ex pend i-tune w as m easured fo r 5 h (100 0- 1 500 h ), and b lo od sam plesw ere taken IS , 3 0 , 45 , 60 , 75 , 90 , 10 5 , 120 , 150 , 180 , 210 , 240 ,an d 3 ( X ) m m afte r the m ea l w as consum ed . D uring the pos tp nan-d ia l m easurem en ts th e sub jec ts w ere a llow ed to w atch ligh t en -te rta inm ent m ov ies , an d to hav e a b reak of a m axim um of 5 m mafte r 2 and 4 h . D ur in g the break the su b jects cou ld sit, w a lkq u ietly , o r g o to the to ile t. T he exac t tim e schedu le w as no tedand repea ted on th e fo llow ing test days . W ater con sum ption du n-in g the te st pe rio d w as al low ed , bu t th e to ta l am oun t c onsum edw as n o ted and repea ted on the secon d tes t d ay .

Im m ed ia te ly b efo re and every 30 m m afte r the m eal, q uestion -nam es to assess h unger, satie ty , fu llness , and p rospective con-sum ption w ere f illed ou t by each sub jec t. R a tings w ere m ad e o n1 0 0 - m m v isua l ana log ue scales (V A S ) w ith w ords ancho red a t

each en d , expressing the m ost po sitiv e (ie , go od , p leasan t) o r them os t nega tive ra ting s (ie , bad , u np leasan t) (2 3). Im m edia tely a f-ten the tes t m ea ls the pa latab ility , ta ste , a fte rta ste , tex tu re, andvisua l app eal of the tw o tes t m ea ls w ere re corded by the sub jec tsus ing V A S scon es. D ata on ene rgy e xpe nditu re are b ein g p ub -lished sep ara te ly (A T ag lia bue , A R ab en , M L H e ijn en , PD eu senben g . E P asq ua li, A A strup , u npub lished observa tions ,1994) . abora tors ana lyses

B l o o d w as sam pled w itho u t stasis th ro ugh the indw elling an -tec ub ita l c an nula by us ing ice d syringe s. P la sm a gluco se an dlac ta te w ere an a ly zed by standard enzym atic m ethod s (2 4).P lasm a g lycero l w as ana lyzed af te r tn ich lonace tic ac id prec ip i-ta tion , essen tia lly as desc rib ed by C h ern ick (25). B lo od fo r de-te rm ina tion of p lasm a catecho lam ines w as co llec ted in tu bescon ta in ing ethy len e g lyco l-b is(f l-am ino ethy l e ther ) -N.N.N ,N-te traace tic ac id (E G T A ) and g lu ta th ion e. Sam p les w ere im m e-

d ia te ly cen trifu ged fo r 10 m m at 3 00f x g and 4 { 1 76}Cnd thep lasm a stoned a t -80 { 176} Cn til de te rm ina tio n of ca tech o lam in esby a rad ioenzym atic m eth od (26 ). Im m un oreac tive insu lin con -cen tra tions w ere m easured in p lasm a w ith rad io im m u noassayk its pu rchased from N ovo , C o penhagen . G astric inh ib ito ry po ly -pep t ide (G IP) and g lucagon -lik e pep tide - I (G L P- I ) w ere de te r-m ined by rad io im m u noassay on p lasm a ex trac ted w ith ethano las p rev ious ly d esc ribed (27 -29). S am p les fo r th e G IP andG L P - I an alyses cou ld no t be taken from th e Italian su b-jects . T he sta rch ana lyses w ere p erfo rm ed at the D un n C lin icalN utrition C en tre (U nited K ingd om ) by us in g the m etho ds de-sc n ib ed by E ng lys t e t a l (2 1 ) a nd E ng ly st a nd C u m m in gs (22 ).

S ta tis tica l a na ly sesA ll resu lts a re g iv en as m eans SE M . D ata a re p resen ted as

chan ges fro m basa l fas tin g concen tra tions by using the seco ndfas ting b lood sam ple as the b asa l con cen tra tion . R espo nses to thetw o tes t m ea ls w ere co m pared b y param etric ana ly sis o f v ar iance(A N O V A ) for rep ea ted , pa ined m easures w ith tim e and d iet asfac to rs an d w ith sub jec t nes ted in to d ie t. T h e k ey resu lts are p re -sen ted w ith F or t va lues an d degrees o f freedo m . A reas underthe curv es (A U C s) fo r the 5-h m easurem en t period s w ere ca l-cu la ted sep ara te ly fo r each sub jec t as the d iffe rence be tw een thein teg ra ted area of the response curve and the rec tangu la r a read ete rm ined by the basa l va lu es ( ne t respon se). A p aired t testw as used in the com p arisons be tw een tw o m eans o n th e sam esub ject. R egress ion an a ly ses w ere perfo rm ed o n the d iffe rencesbe tw een the S and R m eals ( re fe rred to as S -R ) fo r the 5- h A U C sand m eans fo r the V A S scores, peak /nad ir va lues, and peak /L nad ir va lues. T h is w as done to accoun t fo r th e da ta be in gpa ired . T h e leve l o f sig n ificance w as set a t P < 0 .05 . S ta t g r a phic s so ftw are v ersion 4 .2 (G raph ic S oftw are Sy stem s, Inc .R ockv ille , M D ) an d th e S ta tistica l A na lys is P ackag e ( S A SIns titu te , C any , N C ) w ere u sed in the sta tis tica l ca lcu la tions .

R esults

G luco se a nd la c ta teF ast ing p lasm a g luc ose co nc en trations ave rag ed 4 .9 7 0 .08

m m ol/L b efo re th e S m eal an d 4 .9 1 0 .07 m m olIL be fore th eR m eal (N S ). A s ign ifican t in te rac tio n be tw een d ie t and tim e w asobserved afte r the tes t m ea ls w ith g lucose con cen tra tions inc reas-ing n in e t im e s as m uc h after the S m eal a s af ter the R m eal(L p eak con cen tra tions : 5 , 3 .07 0 .29 m m ol/L after 30 m m ; R ,0 .36 0 .13 m m ol/L after 15 m m ) (F1122151 25. 2 , P < 0.0 00 1)(F ig 1). T he 5- h A U C s averaged 79 .53 27 .26 m m ol min /Lafter the S m ea l an d -42 .02 9 .5 1 m m o l m in lL afte r the Rm eal U , 4 .5 1 P 0 .0015). T he A U C s for th e first 2 h a lsoresu lted in a n eg a tive a rea fo r the R m eal 13.7 6 .71m m ol m in lL ) com pared w ith the S m eal ( 140 .00 21 .53m mol m in /L ) t5 7 . 1 1 , P < 0. 0001) .

P lasm a lac tate con cen trations inc reased af te r bo th tes t m ealsw ith a peak afte r 45 m m for th e S m eal (1 .36 0 .0 9 m m olIL ),and afte r 1 5 m m for th e R m eal ( 1 I 8 0 .1 0 m m o l/L ) ( in te rac -t io n d ie t- ti me : F,,215 , 25. 2 , P < 0.0 00 1) (F ig I). T h e increasefrom basa l w as tw ice as h igh afte r the S (0 .52 0 .06 m m o IIL )than af te r the R m eal (0 .2 4 0.0 6 m m ol/L ) . F ive-ho ur A U C saveraged 6 .9 10 .0 m m ol m in /L after the S m eal and -2 6 .3 8 .9 m m olm in lL afte r the R m eal (t9 3. 70 , P 0 .005) .


4/8

5-h ou r A U Cm m o l { 1 4 9 }m { 1 4 9 }

Pl asm a gl ucosemm o l LE

-2 .0 -1 .0 0 .0 1 .0 2 .0 3 .0 4 .0 5 .0

Plasm a lact at em m o l

, < 0.9 71

5 - ho u r AUCm m o l m m L

.20

.40 R S-2.0 -1 .0 0. 0 1 .0 2 .0 3 .0 4 .0 5 .0

Plasm a insu l inp m o l L

, < OM O

-20 -1 .0 0 .0 1 .0 2 .0 3 .0 4 .0 5 .0 R S

Plasm a G I Pp m o l L

40I

20 ,0 . --H- - {149} - - - {149} - -- 2 . 0 -1.0 0.0 1.0 2.0 3.0 4.0 5.0

Pl asm a G L P-lp m o l L

5 - h o u r AU Cn m m m L

iti(

5 - h o u r AU Cn m m m i {1 49 }

- 2 . 0 -1.0 0.0 1.0 2.0 3.0 4.0 5.0H o u r s after test m ea l

F IG I . C hang e in p lasm a concen tra tions o f g lucose , lac ta te , in su lin ,gas tric in h ib ito ry polype p tide (G IP ), a nd gluca gon -like pep tid e-l (G L P -I ) a fte r a raw pota to s tarch m ea l 154 % res istan t stanch (R S)] (R ) and ap rege la tin ized po ta to sta rch m eal (0% R S ) (5 ) in 1 0 hea lthy , no rm al-w eig ht. m a le subje cts . A ll da ta arc m ean ( S E M ), e xpre ssed a s diffe -rcnccs fro m fas ting concen tra tions . L e f t p an el (A N O V A ): a ll (tim eef fect : P < 0 .00 01 ); p lasm a glu cose (d ie t ef fect : F, , . ,5 , = 25 . 1. P

< 0 .0 001 ; in teraction betw ee n d iet a nd tim e: F,,2 5, = 25 .2 , P< 0 .0001) : p lasm a lac ta te (d ie t effec t: F,,,5, = 36. 7 . P < 0 .0001: inter-a ctio n b etw ee n die t an d tim e : F, ,2 5 , = 25 .2 . P < 0 .0001) : p lasm a in -su lin (d ie t e ffec t: F,,.,5, = 39 .9 , P < 0 .0001 : in te rac tion be tw een d ietand tim e: F,5 ,43, = 15 .1 , P < 0 .00 01) ; p la sm a G IP (is = 5) (d ie t e ffe ct:F, , .5 , = 27 .3 , P < 0 .0001 : in te raction be tw een d ie t and tim e: F,5 .5 3 ,= 14 .6 , P < 0 .0001) : p lasm a G LP-1 (, = 5 ) (in te rac tion be tw e en dietand tim e: F,5..,7, = 4 .7 , P < 0 .0001 . R igh t pane l: a reas un der the cu rves( A U C ) . D < 0. 05 . ** < 0 .01 , * p < 0 .001 .

R E S IST A N T S T A R C H , G L Y C E M IA , A N D S A T IE T Y 547Insulin, GIP , a nd G LP - I

F as ting p lasm a in su lin concen tra tio ns averaged 53 .7 5 .0pm ol/L befo re the S m eal and 75 .4 10 .0 pm olIL befo re the Rm eal (N S ). A s ig n ifican tly d iffe ren t respon se p atte rn w as ob-se rved after th e tw o tes t m ea ls . C o m pared w ith fas ting con cen-trations p lasm a insu lin inc reased by a fac to r o f six to 355 .4 68 .5pm ol/L af ter the S m eal , b u t to o n ly 84 .6 7 .2 pm o llL afte r th eR m eal (in te rac tion d ie t-tim e : F 1 5 1 4 = 15 .14 , P < 0.000 1 ) (F igI ). T he A U C s averaged 19 . 1 4 .0 nm ol . m i n / L afte r the S m ea land -6 .6 I .2 nm ol . m i n / L afte r the R m eal t9 = 4 .90 , P= 0 .0008) . A lso , the A U C s for th e f ir st 2 h resu lted in d iffe rencesbe tw een th e S (18 .4 2 .8 nm olm in /L ) and the R m eals (-1 .4 1 .1 nm olm in /L ) t9 = 5 .34 , P = 0 .0005) .

P lasm a G IP show ed the sam e response pa tte rn as p lasm a in -su lin (F ig I . T hus com p ared w ith fas tin g va lu es , G IP increasedby a fac to r o f 10 afte r the S m ea l w hereas n o increase w as ob-serv ed a fter th e R m e al (in te rac tion d iet-tim e: F 1 5 6 1 = 14 .6 , P< 0 .0001). T he A U C S -R for G IP and lac ta te w ere pos itive lycorrelated r = 0 .91 , P = 0 .03) .

P lasm a G L P -I inc reased sign if ican tly a fte r the S m ea l w h ereasn o changes w ere observed afte r the R m eal (in te rac tio n d iet-tim e :F1 8 . 71 4 .7 , P = 0 .000 2) (F ig I . T h e d if fe rence in the A U C safte r the R and th e S m eals w as n early s ign ifican t O = 2. 56 , P

5-ho ur A U C = 0 .06). T he A U C S -R fo r G L P -l w as nega tive ly corre la tedn m o l m m L w ith G IP r = -0 .94 , P 0 .02) and lac ta te r - - 0.88, P

= 0 .0 52) and L peak S -R for G L P -l nega tiv ely corre la tedw ith G IP r = -0 .91 , P = 0 .03) .

Trig lycerides TG ) a nd g lycero lP lasm a trig ly ceride con cen trations sho w ed a s im ila r resp onse

p a tte rn after the tw o test m ea ls (F ig 2 ). T hu s, T G decreaseds ligh tly a fte r bo th m ea ls w ith a nad ir a fte r 2 .5 h an d re tu rned tob ase lin e a fte r 5 h (tim e effec t, F,9 1 51 = 4 .1 , P < 0 .001) . T ri-g ly cer id e concen tra tio ns w ere s ign ifican tly corre la ted w ith in -su lin for the S -R A U C s r = 0 .65 , P = 0 .04) and the S-RL. peak /L na dir va lue s r = 0 .83 , P = 0 .006) .

A tendency to a d ifferen t respon se pa tte rn fo r p lasm a g lycero lw as fou nd afte r the tw o tes t m eals . T h us p lasm a g lycero l de-c reased afte r th e S m ea l to a n ad ir a fte r I . 5 h w hereas no decreasew as ob served afte r the R m eal (in te rac tion d ie t-tim e : F 15 Js j= 1 . 9 , P = 0 .06) (F ig 2) . A fte r 5 h g ly cero l concen tra tions hadincreased by 59 .3 m m olIL fo r the S m ea l and b y 39 .8 m m o lILfo r the R m eal com pared w ith fas tin g con cen trations . T h e A U C sw ere no t s ign ifican tly d iffe ren t a fte r th e tw o test m eals . T heA U C S -R for g ly cero l w as neg ative ly corre la ted w ith g lucose r = -0.75, P = 0 .0 1), w h ereas peak S -R for g lycero l w aspo sitive ly corre la ted w ith peak S-R tr ig ly ceride r = 0. 72 ,P = 0 .02) .


5/8

m m ol L P lasm a tr ig ly cer id es

-.-- A

-0 .1

tw een afte rtas te and m ean sa tie ty r = 0 .65 , P = 0 .04), hun gerr = -0 .64 , P = 0 .0 49 ). an d fu llnes s r = 0 .62 , P = 0 .054) .

Discus s ionM arked d if fe rences in plasm a con cen tra tion s of su bstra tes a nd

horm ones as w ell as in pa la tab ility an d sa tie ty scores w ere ob -se rve d after the tw o te st m e als . O ve rall , the S m ea l g rea tly sti-m ula ted p lasm a co ncen tra tion s o f g lu cose , insu lin . and gas tro -in te stinal h orm o nes w hereas n o or on ly a m od est st im u lat ion ofthese indexes w as observed afte r the R m eal.

P lasm a g lucose inc reased n ine tim es m o re afte r th e S m ea l thanafte r the R m eal. T h is w as no t en tire ly unexpec ted on the bas iso f the an aly tica l da ta on d igestib ility o f the tw o tes t sta rches.H ow ever, the d iffe rences in p ostp rand ia l g lycem ia w ere som e-w ha t la rger th an w ould h e ex pected from th e clin ica l ana lyses.T hus , the ratio be tw e en the p eak g luc ose respon se af ter the Rand S m eal w as I :9 (0 .361 3 .0 7 m m oIlL ), w hereas the ra tio b e-tw een th e am ount o f read ily d igestib le ca rboh ydra te w as 1 :5( I I . 4154 .4 g) (T ab le 3 ). T he reason fo r th is d iffe rence is no t rca -d ily ap paren t bu t m ay be con ne cted w ith d ifferenc es in the s ti-m ula tio n of gas tro in tes tina l facto rs and insu lin af te r th e tw omeals .

D esp ite the m inor in crease in p lasm a g lu co se afte r the R m eal,the in crease in p lasm a lac ta te concen tra tion am ounted to 50 % ofthe in crease a fte r the S m ea l. T h is m ay be due to the fruc tosepresen t in the tes t m ea l (4 .4 g) o r to a part o f th e g lucose be ingconv erted to lac ta te v ia non ox ida tive p athw ays in the sp lanchn icr e g i o n ( 3 0 ) .

A lso , p lasm a in su l in increased m ark ed ly af ter the S c om paredw ith th e R m eal , p ro du cin g a rat io be tw e en sp eak va lue s of 30 : 1( I 82 /6 p m o IIL ). T h is d ifferenc e in insu lin resp on se is fa r g re ater

P lasm a n or ep in ep hrm epg {149}l -.-- A 5-hou r A U C

4 0 - .&-s n g m i n L 2:H*/ i : : , Ac >

- 2 . 0 - 1 . 0 0 . 0 1 . 0 2 . 0 3 . 0 4 . 0 5 . 0 R S

P la sm a ep in ep hr in ep g {149}l . 5-hour A UC

5-hour A UCi nmo l mm L

s

-2 .0 -1 .0 0 .0 1 .0 2 .0 3 .0 4 .0 5 .0 A S

P la sm a gly cer o lI 5-ho ur A U C n o l L

1 0 0 i n mo l mm L8

50 r_ ::: ;_2 :_ : i : 6i0 4

. . . . TT- 2 . 0 - 1 .0 0 .0 1 .0 2 .0 3 .0 4 .0 5 .0 R S

H ours a fte r tes t m ealF IG 2 . C hange in p lasm a c onc e ntr a t ions of trig lyc eride s and gly ce ro l

a fter a raw p otato s ta rc h m e al (5 4% res is tan t s tarc h IR S I) (R ) and ap re ge latin iz ed potato s tarc h m e al 0 7 R S ) (S ) in 1 0 h eal thy , no rm al-w eig ht. m a le su bjec ts. A ll data a re m e an ( S E M ) e xpre ssed as ch ang esf rom fa stin g co nce ntra tio ns. L eft pane l (A N O V A ): a ll (tim e effect: P


6/8

iT S E M . R , raw p otato s tarch m eal: S . p reg elatin iz ed p otato s tarchm e a l .

2 4 S ignif ic antly dif f erentf rom R (unp aired ltest, df= 9) : O .O ( ) 7 .3P 0.02. 4P O.OO( ) 4 .

5. 02. 50. 0

-2.5 1

H ow hungr y ar e youI h a ve n e ve r b e e n m o re h un gry

5 .0 I

5 6I a m n o t h u n g ry a t a t

H ow m uch do you t h in k you can eatNo th in g a t a ll_ j1 A lot

H o u rs af te r te s t m eal

1 0 1 2 3 4 5 6I a m c o mp le te ly e mp ty

T ot al ly fu ll5. 02. 50. 0

2 .5

H ow fu l l do you f eel

I 1 0 1 2 3 4 5 6

N ot a t a M h itH o u rs af te r te st m eal

FIG 4 . S u b jec tiv e satie ty sco res af te r a raw p o tato s tarch m eal (5 4 re sis tan t starch IR S I ) (R ) an d a p reg e latin iz edpotato starch m eal (0 R S ) (5) in 1 0 he althy . n orm al-w e igh t. m ale su bje cts. A ll d ata are m e an (S E M ). ex pre sse d asdif f erenc es f rom f astin g v alue s. A N O V A : all (tim e ef f ect: F,,,.,951. P < 0.000 1): satiety (top righ t): d ie t ef f e ct: F ,,,,, 4.94, P 0 .0 4 : in te rac tio n b e tw een d ie t an d tim e: F ,,,,,,, 1.95, P 0 .0 3 ). Fu lln ess (rig h t b o tto m ): in te rac tio nb e tw een d iet an d tim e: F ,,,,95, 2.30. P 0.012) .

R E S IS T A N T S T A R C H , G L Y C E M IA . A N D S A T IE T Y 5 4 9th an w o u ld b e ex p ected f ro m th e d if f e ren ce in g lu co se lo ad an dg lu co se resp o n se af te r th e tw o m eals . In try in g to ex p lain th is itm ay b e u se f u l to lo o k at th e ch an g es in g as tro in tes tin al h o rm o n es .N ot on ly did the R m eal re sult in a m u ch low er blo od g luc oseresp o n se co m p ared w ith th e S m eal, b u t at th e sam e tim e th ereseem ed to b e n o e f f ec t o n e ith er th e p ro x im al (G IP) o n d is tal(G L P- I ) incret in s in co n trast w ith th e in crease in th ese h o rm o n esaf te r th e S m eal. B o th G IP an d G L P- I are k n o w n to b e p o ten ts tim u lato rs o f in su lin secre tio n (3 1 . T h e larg e d if f e ren ce b e -tw e en the m eals in th ese h orm on es m ay th ere f ore ex plain th ed if f e ren ce -b ey o n d w h at can b e ex p ec ted f ro m th e g lu co se -s tim u lated in su lin secre tio n -in in su lin re sp o n se b etw een th etw o m eals .

T h e su b jectiv e f ee lin g s o f satie ty an d f u lln ess w ere also s ig -n if ican tly in f lu en ced b y th e d ig estib ility o f th e starch . T h u s , th ep reg elatin iz ed and f u lly d ige stib le potato starch m eal resu lte d ing reate r f ee lin g s o f satie ty an d f u lln ess co m p ared w ith th e n es is-tan t an d slo w ly d ig estib le starch m eal. A lread y , 1 -1 .5 h af te r th eraw p o tato s tarch m eal th e su b jec tiv e sco n es w ere b ack to f as tin gco n cen tratio n s, w h ereas th e satiatin g p o w er o f th e S m eal las ted2 . 5 3 h p o s t p r a n d i a l l y .

It h as p rev io u sly b een stated th at ch an g es in satiety af te r acarb o h y d rate lo ad m ay b e m ed iated th ro u g h an e f f ec t o f p lasm ag lu co se o n h ep atic g ly co g en co n cen tratio n o n sp ec if ic g lu co sen -s itiv e cells in the brain ( 32 ) an d s tu d ie s h av e d em o n s trated asatiatin g e f f ec t o f carb o h y d rate p er se ( 1 4 ). W e f o u n d n o s ig n i-f ican t co rre latio n s b e tw een th e ch an g es in p lasm a g lu co se an d inth e f o u r satie ty sco res. In a recen t stu d y f ro m o u r d ep artm en t inw h ich a h ig h - an d a lo w -f ib er m eal w ere g iv en to 1 0 h ealth ysu b jec ts, th e A U C s f o r g lu co se an d satie ty sco res w ere n o t sig -n if ican tly co rre lated e ith er ( I 8 ). In th e p resen t s tu d y d if f e ren cesin hu nge r rating s be tw een the tw o m eals w ere in f act co rrelate dw ith d if f e ren ces in L p eak lac tate co n cen tratio n , w h ich m ay re -

T A B L E 4S ubjectiv e ev aluation of th e test m e al

R SV isual appe al (0 : g ood . 10: b ad )T aste (0 : p leasan t. 1 0 : u n p leasan t)A f tertaste (0: n one , 10 : m uc h)T ex tu re (0 : f irm . 1 0 : lo o se)Palatabil ity (0: goo d. 1 0: bad)

5. 3 0. 76. 0 t) .85.4 I . I7 .7 0 .87 .4 0 .6

8.1 0 .57.2 0 .87 .0 0 .73 .4 O W8 .2 0 .6

f lect th e rate o f n o n o x id ativ e g lu co se d isp o sal. In th e p resen ts tu d y carb o h y d rate w as th e o n ly en erg y so u rce in th e te st m eals ,b u t b ecau se an in creased en erg y lo ad in a m eal h as also b eensh o w n to resu lt in an in creased satiatin g p o w er o f th e m eal (3 3 ),it m ay be that th e dif f erenc es in satiety se nsatio ns w e re d ue to ad if f e ren t n et en erg y lo ad in th e m eals . A f irm er co n c lu s io n m ay ,h o w ev er, req u ire m o re d irec t m easu rem en ts o f g ly co g en s to reso r g lu co se d isp o sal in o rd er to ex p lain ch an g es in su b jec tiv e sa-tie ty sco n es .

G as tro in tes tin al h o rm o n es h av e p rev io u sly b een co n n ec tedw ith satiety and ob es ity (34 , 35 ). In the prese nt stud y so m e co n-re latio n s w ere f o u n d b e tw een G IP an d satie ty sco n es b u t n o t b e -tw een G L P-l an d satie ty . T h e co rre latio n s f o u n d in d icate th atG IP m ay d ecrease satie ty an d in crease h u n g er, w h ich is in co n -trast w ith th e p rev io u sly su g g ested satiatin g p o w er o f G IP. In th ep resen t s tu d y , h o w ev er, d ata o n G IP w ere o n ly av ailab le f o r f iv esu b jec ts . T h e co rre latio n s m u st th ere f o re b e co n sid ered w ith

H ow sat i sf i ed d o you f eelI c a n n o t e a t a no th er b ite

. R


7/8

55 0 R A B E N E T A Lsom e cau tio n , an d need to be conf irm ed befo re any fu rthe r con-c lus io ns can be m ade .

B ecause th e v o lum e of th e tes t m ea ls and the tim e sp en t oncon sum ing the m ea ls on the 2 tes t days w ere sim ilar , these fac to rscanno t have in flu enced the sa tia tin g po w er of th e tw o m eals .H o w ever, th e ge la tin iza tion of the S m eal and the fact tha t it w asa so lid m ea l m ay have resu lted in a red uc tio n in g as tric em pty ingra te (3 6) and in in creased fee lin gs of sa tie ty and fu llness a fte rth is m eal com p ared w ith the liqu id R m eal. A lso , th e d ifferen tra tings of taste , v isua l appea l, and tex tu re m ay have in fluencedsatie ty and fu lln ess ra tin gs. In fac t, p ositive corre la tion s w erefoun d be tw een afte rtas te an d sa tie ty and fu llness. T h is partly con-firm s the prev ious stud y by H ill e t a t (2 3) in w hich increasedhun ger w as found w ith inc reased p re fe ren ce (pa latab ility ) o f am ea l. H ow ever, in th e ir stud y there w as no effec t on fu llnessra tin gs of the m ea l p re fe rence (23).

T he effec t o f R S on sa tie ty sho w n in the p resen t s tudy m ay becons idered a nega tive e ffec t o f R S w ith regard to appe tite regu-lation an d perhaps a lso w eig h t con tro l. R ep lacem en t o f d iges tib lesta rch w ith no nd igestib le sta rch in the d ie t m ay pose a risk ofinc reasing the overa ll fa t- to -carb ohyd ra te ra tio in th e d ie t. B e-cause a h igh-fa t, low -carbohy dra te d ie t has been sho w n to in -c rease energy in take and body w eig h t under ad lib itum con d i-tio ns, w h ereas a low -fa t, h igh-ca rboh ydra te d ie t h as been sh ow nto decrease spon taneo us food in take and resu lt in unexp ec tedw eigh t reduc tio ns (37-39), a rep lacem en t o f d ig es tib le sta rchw ith R S m ay be un desirab le in long -term w eig h t regu la tion .W hether R S in a m ixed m ea l h as the sam e effect o n appe tite asin th e p resen t stud y rem ains , how ever, to be e luc ida ted . M o re-over, it is no t p oss ib le to pred ic t the e ffec t on 2 4-h appe tite sen -sa tions from the presen t 5 -h resu lts . T he R S not d igested in thesm all in tes tin e w ill at a la te r s tag e, ie , 7 - 12 h af te r ingestion , bed igested and fe rm en ted by bac te ria in the la rg e in testine , resu ltingin th e p rod uct ion an d up tak e o f shor t-ch ain fa tty ac ids (5 , 6 , 40).W hether th is m ay co n tribu te to in c reas ing sa tie ty rem ains to bee luc ida ted .

In co nc lusion , th e in take of R S resu lted in s ig n ifican tly low erpostp ran d ia l p lasm a g lucose , lac ta te , in su lin , G IP , and G L P -lrespon ses and in a reduc tio n o f the sa tiating po w er co m paredw ith d igestib le s tarch . T he ch anges in p lasm a g lucose w ere no tco rrela te d w ith satie ty or fu l lne ss, bu t co rre la tions w e re fo undbe tw een G IP an d sa tiety scores . D iffe rences in tex tu re and pa l-atab ility o f the test m ea ls m ay a lso h ave in fluenced the sub jec tiv esa tie ty ra ting s. F urthe r stud ies a re needed to c la rify th e effec t o f in a m ixed m ea l, bo th acu te ly and long te rm . U

W e grate fu lly thank L ene M ellem , K jas D cnsen , the labora to ry tech-nicians B en te K n ap . Inge T im m erm ann , John L ind , B en te M ath iase n .an d K aren K lausen : the d ie titians C laud ia T rcn tan i and C ate rina M am in i:an d lng cr-L ise G r {2 48}n fe ld t n d her k itchen s ta ff fo r expert techn ica l ass is -tance as w ell as G re the T h orbek fo r consu ltive assis tance .

e fe r ence sI . D ahlqv ist A , B o rg trO m B . D iges tion and a bsorp tion o f d isacc har ide s

in man. Biocheml 1961 :81 :411-8 .2. E nglys t H N , W igg ins H S , C um m ing s IH . D ete rm in ation of the n on-

s tarch po lysaccharides in p lan t foods by gas-liq u id chro m atog raphyof con stitu en t su gars as a ld ito l ace ta tes. A na lys t 1982 :1 07 :307-18 .

3 . S tep hen A M . H addad A C . P h illip s SF . P assage o f ca rbohy dra te in tothe co lon . D irect m easurem en ts in h um ans. G astroen teno log y1983:85 :589-95 .

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resistant starch - the effect on postprandial glycemia, hormonal response, and satiety

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