Abstract

Inqubekelaphambili enkulu yocwaningo lwama-carbohydrate ekudleni kwezingulube kanye nempilo ukuhlukaniswa okucacile kwe-carbohydrate, okungasekelwe kuphela ekwakhekeni kwamakhemikhali, kodwa futhi ngokusekelwe ezicini zayo zomzimba. Ngaphezu kokuba umthombo wamandla oyinhloko, izinhlobo ezahlukene kanye nezinhlaka zama-carbohydrate zizuzisa ekondleni kanye nemisebenzi yezempilo yezingulube. Bahileleke ekukhuthazeni ukukhula nokusebenza kwamathumbu ezingulube, ukulawula umphakathi wamathumbu e-microbial wamathumbu, nokulawula i-metabolism ye-lipids ne-glucose. Indlela eyisisekelo ye-carbohydrate itholakala ngama-metabolites ayo (ama-chain chain fatty acids [SCFAs]) futhi ikakhulukazi nge-scfas-gpr43 / 41-pyy / GLP1, SCFAs amp / atp-ampk kanye ne-scfas-ampk-g6pase / PEPCK izindlela zokulawula amafutha kanye ne-glucose metabolism. Ucwaningo olusha luhlole inhlanganisela efanele yezinhlobo nezinhlaka ezahlukene zamacarbohydrates, ezingathuthukisa ukusebenza kokukhula kanye nokugayeka kokudla okunomsoco, kukhuthaze ukusebenza kwamathumbu, futhi kwandise inala yamagciwane akhiqiza i-butyrate ezingulubeni. Sekukonke, ubufakazi obuqand' ikhanda busekela umbono wokuthi ama-carbohydrate adlala indima ebalulekile ekusebenzeni okunempilo kanye nempilo yezingulube. Ngaphezu kwalokho, ukuzimisela kokwakheka kwe-carbohydrate kuzoba nenani lethiyori nelisebenzayo ekuthuthukisweni kobuchwepheshe bebhalansi ye-carbohydrate ezingulubeni.

1. Isandulelo

Ama-polymeric carbohydrates, isitashi kanye ne-nonstarch polysaccharides (NSP) yizingxenye eziyinhloko zokudla kanye nemithombo yamandla eyinhloko ezingulubeni, ezibalelwa ku-60% - 70% wesamba samandla esiwadlayo (Bach Knudsen). Kuyaphawuleka ukuthi izinhlobonhlobo kanye nokwakheka kwama-carbohydrate kuyinkimbinkimbi kakhulu, enemiphumela ehlukene ezingulubeni. Ucwaningo lwangaphambilini lubonise ukuthi ukondla ngesitashi ngesilinganiso se-amylose eya ku-amylose (AM / AP) ehlukile kunempendulo ecacile yomzimba ekusebenzeni kokukhula kwezingulube (Doti et al., 2014; Vicente et al., 2008). I-fiber yokudla, ikakhulukazi eyakhiwe i-NSP, kukholakala ukuthi yehlisa ukusetshenziswa kwezakhi kanye nenani lamandla ezilwane ezi-monogastric (NOBLET kanye ne-le, 2001). Kodwa-ke, ukudla okune-fiber akuthikameze ukukhula kwamaphuphu (Han & Lee, 2005). Ubufakazi obuningi bubonisa ukuthi i-fiber yokudla ithuthukisa i-morphology yamathumbu kanye nomsebenzi wokuvimbela ama-piglets, futhi inciphisa izehlakalo zohudo (Chen et al., 2015; Lndberg, 2014; Wu et al., 2018). Ngakho-ke, kuyaphuthuma ukufunda ukuthi ungasebenzisa kanjani ngempumelelo ama-carbohydrate ayinkimbinkimbi ekudleni, ikakhulukazi ukudla okunothe nge-fiber. Izici zesakhiwo kanye ne-taxonomic yama-carbohydrates kanye nemisebenzi yawo yokudla okunempilo kanye nezempilo ezingulubeni kufanele kuchazwe futhi kubhekwe ekwakhiweni kokuphakelayo. I-NSP kanye nesitashi esingagayeki (RS) zingama-carbohydrate angagayeki ayinhloko (wey et al., 2011), kuyilapho i-microbiota yamathumbu ivubela ama-carbohydrate angagayeki abe ama-short chain fatty acids (SCFAs); Turnbaugh et al., 2006). Ngaphezu kwalokho, amanye ama-oligosaccharides nama-polysaccharides abhekwa njengama-probiotics ezilwane, angasetshenziswa ukukhuthaza ingxenye ye-Lactobacillus ne-Bifidobacterium emathunjini (Mikkelsen et al., 2004; Mø LBAK et al., 2007; Wellock et al., 2008). Ukwengezwa kwe-Oligosaccharide kuye kwabikwa ukuthuthukisa ukwakheka kwe-microbiota yamathumbu (de Lange et al., 2010). Ukuze kuncishiswe ukusetshenziswa kwabagqugquzeli bokukhula kwe-antimicrobial ekukhiqizeni izingulube, kubalulekile ukuthola ezinye izindlela zokuzuza impilo enhle yezilwane. Kukhona ithuba lokwengeza izinhlobonhlobo eziningi zamacarbohydrate ekudleni kwezingulube. Ubufakazi obuningi bubonisa ukuthi inhlanganisela efanele yesitashi, i-NSP ne-MOS ingakhuthaza ukusebenza kokukhula kanye nokugayeka kokudla okunomsoco, kwandise inani lamabhaktheriya akhiqiza i-butyrate, futhi kuthuthukise i-lipid metabolism yezingulube ezilunyuliwe ngokwezinga elithile (Zhou, Chen, et al., 2020; Zhou, Yu, et al., 2020). Ngakho-ke, inhloso yaleli phepha ukubuyekeza ucwaningo lwamanje ngendima ebalulekile ye-carbohydrate ekukhuthazeni ukusebenza kokukhula nokusebenza kwamathumbu, ukulawula umphakathi wamathumbu emathunjini kanye nempilo ye-metabolic, kanye nokuhlola inhlanganisela ye-carbohydrate yezingulube.

2. Ukuhlukaniswa kwama-carbohydrate

Ama-carbohydrate okudla angahlukaniswa ngokwesayizi yawo yamangqamuzana, idigri ye-polymerization (DP), uhlobo lokuxhuma (a noma b) kanye nokwakheka kwama-monomers ngamanye (Cummings, Stephen, 2007). Kuyaphawuleka ukuthi ukuhlukaniswa okuyinhloko kwama-carbohydrate kusekelwe ku-DP yabo, njenge-monosaccharides noma i-disaccharides (DP, 1-2), i-oligosaccharides (DP, 3-9) kanye ne-polysaccharides (DP, ≥ 10), eyakhiwe isitashi, i-NSP kanye nezibopho ze-glycosidic, i-Cummings, i-Englyt, i-7; 2007; Ithebula 1). Ukuhlaziywa kwamakhemikhali kuyadingeka ukuze uqonde imiphumela yokuphila kanye nempilo yama-carbohydrate. Ngokuhlonza amakhemikhali amaningi ama-carbohydrates, kungenzeka ukuwahlanganisa ngokwemiphumela yawo yezempilo neyomzimba futhi uwafake ohlelweni lokuhlukanisa lulonke (englyst et al., 2007). Ama-carbohydrate (ama-monosaccharides, ama-disaccharides, kanye nesitashi esiningi) angagaywa ama-enzyme abamba futhi amuncwa emathunjini amancane achazwa njengama-carbohydrates agayekayo noma atholakalayo (Cummings, Stephen, 2007). Ama-carbohydrate amelana nokugayeka kwamathumbu emathunjini, noma amuncwa kabi futhi afakwe i-metabolized, kodwa angase acekelwe phansi ukuvutshelwa kwe-microbial abhekwa njengama-carbohydrates amelana, njengama-NSP amaningi, ama-oligosaccharides angenakudliwa kanye ne-RS. Empeleni, ama-carbohydrate amelana achazwa njengangagayeki noma angasebenziseki, kodwa anikeze incazelo enembe kakhulu yokuhlukaniswa kwama-carbohydrate (englyst et al., 2007).

3.1 ukukhula kokusebenza

Isitashi sakhiwe izinhlobo ezimbili zama-polysaccharides. I-Amylose (AM) iwuhlobo lwesitashi esiwumugqa α (1-4) i-dextran exhunyiwe, i-amylopectin (AP) iyi-dextran exhunywe ku-α（ 1-4), equkethe cishe u-5% dextran α(1-6) ukuze yakhe i-molecule enegatsha (umhloli et al., 200). Ngenxa yokucushwa kwe-molecular eyahlukene kanye nezakhiwo, i-AP isitashi esicebile kulula ukugayeka, kuyilapho isitashi esicebile akulula ukusigayeka (Singh et al., 2010). Ucwaningo lwangaphambilini lubonise ukuthi ukondla isitashi ngezilinganiso ezihlukene ze-AM / AP kunezimpendulo ezibalulekile zomzimba ekusebenzeni kokukhula kwezingulube (Doti et al., 2014; Vicente et al., 2008). Ukudla okuphakelayo kanye nokusebenza kahle kokuphakelayo kwezingulube ezilunyuliwe kwehle ngokwanda kwe-AM (regmi et al., 2011). Kodwa-ke, ubufakazi obusafufusa bubika ukuthi ukudla okunezinga eliphakeme kakhulu kwenyusa inzuzo yansuku zonke kanye nokusebenza kahle kokudla kwezingulube ezikhulayo (Li et al., 2017; Wang et al., 2019). Ngaphezu kwalokho, abanye ososayensi babike ukuthi ukuphakela okuhlukile kwe-AM / AP kwesitashi akuzange kuthinte ukukhula kwezingulube ezilunyuliwe (Gao et al., 2020A; Yang et al., 2015), kuyilapho ukudla okuphezulu kwe-AP kukhuphule ukugayeka kokudla kwezingulube ezilunyuliwe (Gao et al., 2020A). I-fiber yokudla iyingxenye encane yokudla evela ezitshalweni. Inkinga enkulu ukuthi i-fiber yokudla ephezulu ihlotshaniswa nokusetshenziswa kwezakhi eziphansi kanye nenani eliphansi lamandla (noble & Le, 2001). Ngokuphambene nalokho, ukudla okusesilinganisweni kwe-fiber akuzange kuthinte ukukhula kwezingulube ezilunyuliwe (uHan & Lee, 2005; Zhang et al., 2013). Imiphumela ye-fiber yokudla ekusetshenzisweni komsoco kanye nenani lamandla eliphelele lithintwa izici ze-fiber, futhi imithombo ye-fiber ehlukene ingase ihluke kakhulu (lndber, 2014). Ezingulubeni ezilunyuliwe, ukuxhaswa nge-pea fiber kwakunezinga eliphezulu lokuguqulwa kokuphakelayo kunokondla i-corn fiber, i-soya fiber kanye ne-wheat bran fiber (Chen et al., 2014). Ngokufanayo, amazinyane ezingulube alunyuliwe aphathwe nge-corn bran kanye ne-bran kakolweni abonise ukusebenza kahle kokuphakelayo kanye nokuzuza kwesisindo kunalawo aphathwe ngobhontshisi wesoya (Zhao et al., 2018). Kuyathakazelisa ukuthi awukho umehluko ekusebenzeni kokukhula phakathi kweqembu le-fiber bran kakolweni neqembu le-inulin (Hu et al., 2020). Ukwengeza, uma kuqhathaniswa namaphuphu aseqenjini le-cellulose kanye neqembu le-xylan, ukwesekwa kwakusebenza kangcono i-β-Glucan ikhubaza ukusebenza kokukhula kwezingulube (Wu et al., 2018). Ama-Oligosaccharides ama-carbohydrate anesisindo esiphansi samangqamuzana, aphakathi phakathi kukashukela nama-polysaccharides (voragen, 1998). Zinezinto ezibalulekile ze-physiological and physicochemical, kufaka phakathi inani eliphansi le-calorific kanye nokukhuthaza ukukhula kwamagciwane anenzuzo, ngakho-ke angasetshenziswa njengama-probiotics okudla (Bauer et al., 2006; Mussatto kanye ne-mancilha, i-2007). Ukwengezwa kwe-chitosan oligosaccharide (COS) kungathuthukisa ukugayeka kokudla okunomsoco, kunciphise ukwenzeka kwesifo sohudo futhi kuthuthukise i-morphology yamathumbu, ngaleyo ndlela kuthuthukise ukusebenza kokukhula kwezingulube ezilunyuliwe (Zhou et al., 2012). Ngaphezu kwalokho, ukudla okunezelwa nge-cos kungathuthukisa ukusebenza kokuzala kwezingulube (inani lezingulube eziphilayo) (Cheng et al., 2015; Wan et al., 2017) kanye nokusebenza kokukhula kwezingulube ezikhulayo (wontae et al., 2008). Ukwengezwa kwe-MOS ne-fructooligosaccharide kungabuye kuthuthukise ukusebenza kwezingulube (Che et al., 2013; Duan et al., 2016; Wang et al., 2010; Wenner et al., 2013). Le mibiko ikhombisa ukuthi ama-carbohydrate ahlukahlukene anemiphumela ehlukile ekusebenzeni kokukhula kwezingulube (ithebula 2a).

3.2 ukusebenza kwamathumbu

Isitashi esiphezulu se-am/ap singathuthukisa impilo yamathumbu (tribyriningavikela ingulube) ngokuthuthukisa i-morphology yamathumbu kanye nokulawula ukusebenza kwamathumbu okuhlobene nokuvezwa kofuzo ezingulubeni ezilunyulwayo (Han et al., 2012; Xiang et al., 2011). Isilinganiso sobude be-villi nobude be-villi nokujula kwe-recess ye-ileum ne-jejunum sasiphezulu uma siphakelwa ngokudla okuphezulu kwe-am, futhi inani eliphelele lesilinganiso se-apoptosis lamathumbu amancane laliphansi. Ngasikhathi sinye, iphinde yandisa ukubonakaliswa kokuvimbela izakhi zofuzo ku-duodenum nejejunum, kuyilapho eqenjini eliphakeme le-AP, imisebenzi ye-sucrose ne-maltase ku-jejunum yezingulube ezilunyuliwe yanda (Gao et al., 2020b). Ngokufanayo, umsebenzi wangaphambilini wathola ukuthi ukudla okucebile kwehlise i-pH kanye nokudla okucebile kwe-AP kwandise inani eliphelele lamagciwane ku-caecum yezingulube ezilunyuliwe (Gao et al., 2020A). I-fiber yokudla iyisici esibalulekile esithinta ukuthuthukiswa kwamathumbu kanye nokusebenza kwezingulube. Ubufakazi obuqoqiwe bubonisa ukuthi i-fiber yokudla ithuthukisa i-morphology yamathumbu kanye nomsebenzi wesithiyo wezingulube ezilunyuliwe, futhi inciphisa izehlakalo zohudo (Chen et al., 2015; Lndber, 2014; Wu et al., 2018). Ukushoda kwe-fiber yokudla kukhulisa ukuthambekela kwamagciwane futhi kukhubaza umsebenzi wokuvinjwa kwe-colon mucosa (Desai et al., 2016), ngenkathi ukudla ngokudla kwe-fiber engancibiliki kakhulu kungavimbela amagciwane ngokwandisa ubude be-villi ezingulubeni (hedemann et al., 2006). Izinhlobo ezahlukene zemicu zinemiphumela ehlukene emsebenzini we-colon kanye ne-ileum barrier. I-wheat bran ne-pea fibers ithuthukisa ukusebenza kwesithiyo samathumbu ngokulawula ukubonakaliswa kofuzo lwe-TLR2 nokuthuthukisa imiphakathi yamathumbu amancane uma kuqhathaniswa nemicu yommbila nesoya (Chen et al., 2015). Ukungenisa isikhathi eside kwe-pea fiber kungalawula i-metabolism ehlobene nezakhi zofuzo noma ukubonakaliswa kwamaprotheni, ngaleyo ndlela kuthuthukise umgoqo we-colon kanye nomsebenzi wokuzivikela komzimba (Che et al., 2014). I-Inulin ekudleni ingagwema ukuphazamiseka kwamathumbu ezingulubeni ezilunyuliwe ngokukhuphula ukuqina kwamathumbu (Awad et al., 2013). Kuyaphawuleka ukuthi inhlanganisela ye-soluble (i-inulin) ne-fiber enganyibiliki (i-cellulose) iphumelela kakhulu kunoyedwa, engathuthukisa ukumuncwa okunomsoco kanye nomsebenzi wokuvimbela emathunjini ezingulubeni ezilunyuliwe (Chen et al., 2019). Umphumela we-fiber yokudla ku-mucosa yamathumbu uncike ezingxenyeni zabo. Ucwaningo lwangaphambilini lwathola ukuthi i-xylan ikhuthaza umsebenzi wokuvinjelwa kwamathumbu, kanye nezinguquko ku-spectrum ye-bacterial spectrum kanye ne-metabolites, futhi i-glucan ikhuthaza umsebenzi wokuvinjelwa kwamathumbu emathunjini kanye nempilo ye-mucosal, kodwa ukuxhaswa kwe-cellulose akuzange kubonise imiphumela efanayo ezingulubeni zokulunyulwa (Wu et al., 2018). Ama-oligosaccharides angasetshenziswa njengemithombo yekhabhoni yama-microorganisms asemathunjini aphezulu esikhundleni sokugaywa futhi asetshenziswe. Ukwengezwa kwe-Fructose kungandisa ukushuba kwe-mucosa emathunjini, ukukhiqizwa kwe-asidi ye-butyric, inani lamaseli aphindaphindiwe kanye nokwanda kwamangqamuzana e-epithelial emathunjini ezingulube ezilunyuliwe (Tsukahara et al., 2003). I-Pectin oligosaccharides ingathuthukisa umsebenzi wokuvimbela emathunjini futhi inciphise ukulimala kwamathumbu okubangelwa i-rotavirus kuma-piglets (Mao et al., 2017). Ngaphezu kwalokho, kutholwe ukuthi i-cos ingakhuthaza kakhulu ukukhula kwe-mucosa yamathumbu futhi ikhulise kakhulu inkulumo yokuvimbela izakhi zofuzo kuma-piglets (WAN, Jiang, et al. ngendlela ebanzi, lezi zikhomba ukuthi izinhlobo ezahlukene ze-carbohydrate zingathuthukisa ukusebenza kwamathumbu emathunjini (ithebula 2b).

Isifinyezo kanye Nethemba

I-carbohydrate ingumthombo wamandla oyinhloko wezingulube, ezakhiwe ngama-monosaccharides ahlukahlukene, ama-disaccharides, ama-oligosaccharides nama-polysaccharides. Imigomo esekelwe ezicini zomzimba isiza ukugxila emisebenzini yezempilo engaba khona yama-carbohydrate futhi ithuthukise ukunemba kokuhlukaniswa kwama-carbohydrate. Izakhiwo nezinhlobo ezahlukene zama-carbohydrate zinemiphumela ehlukene ekugcineni ukusebenza kokukhula, ukukhuthaza ukusebenza kwamathumbu kanye nokulinganisela kwe-microbial, kanye nokulawula i-lipid ne-glucose metabolism. Indlela engenzeka yokulawulwa kwe-carbohydrate ye-lipid ne-glucose metabolism isuselwa kuma-metabolites awo (ama-SCFA), avutshelwa yi-microbiota yamathumbu. Ngokukhethekile, ama-carbohydrate ekudleni angalawula i-glucose metabolism ngokusebenzisa i-scfas-gpr43 / 41-glp1 / PYY kanye ne-ampk-g6pase / PEPCK izindlela, futhi ilawule i-lipid metabolism ngokusebenzisa i-scfas-gpr43 / 41 kanye ne-amp / atp-ampk izindlela. Ukwengeza, lapho izinhlobo ezahlukene zama-carbohydrate zihlangene kahle kakhulu, ukukhula nokusebenza kwezempilo kwezingulube kungase kuthuthukiswe.

Kuyaqapheleka ukuthi imisebenzi engaba khona ye-carbohydrate kumaprotheni kanye ne-gene expression kanye nokulawulwa kwe-metabolic izotholwa ngokusebenzisa ama-proteomics asebenzayo aphezulu, ama-genomics kanye nezindlela ze-metabonomics. Okokugcina, ukuhlolwa kwezinhlanganisela ezahlukene zama-carbohydrate kuyisidingo socwaningo lwezinhlobonhlobo zokudla okunama-carbohydrate ekukhiqizeni izingulube.

Umthombo: Ijenali Yesayensi Yezilwane