Epigenetic Abstract:
Ukunyuka kwenyameko yokunyanya ukugqithisa kunye nokuxhatshazwa okunxulumene nayo yinkinga enkulu yempilo yoluntu. Nangona i-genetic ngokungathandabuzekiyo idlala indima ekunqumeni ukukhutshwa komzimba ngamnye kunye nokukhuluphala, ukuhlukahluka kofuzo oluchongiweyo kuchaza kuphela inxalenye yenguqu. Oku kuye kwakhokelela ekukhuliseni umdla ekuqondeni inxaxheba enkulu ye-epigenetics njengomlamli we-gene-environment interactions ekugxilweni kokunyameka nokuxhatshazwa kwayo. Ubungqina bokuqala bokuxhasa indima ye-epigenetics ekunyanyeni kunye nohlobo lwe-2 yeswekile yesifo sikashukela (T2DM) lubonelelwe ngokubanzi kwizifundo zezilwanyana, ezichaze utshintsho lwe-epigenetic kwiimpawu eziphambili ze-metabolic elandela ukunyusa kwamafutha amaninzi kunye nokwahlukana kwe-epigenetic phakathi kwezilwanyana ezixhambileyo kunye nezilwanyana izifundo zabantu ezibonisa utshintsho lwe-epigenetic ekutyebeni kunye ne-T2DM yegciwane lomviwa kwi-obese / abantu besifo sikashukela. Ngoku kutshanje, ukuqhubela phambili kwiindlela ze-epigenetic kunye neendleko ezincitshisiweyo zophando ze-epigenome-wide width (EWAS) zikhokelela ekunyuseni ngokukhawuleza kwezifundo kubantu. Ezi zifundo ziye zachaza ulwahlulo lwe-epigenetic phakathi kwabantu abadala nabangama-T2DM kunye nokulawula okunempilo kunye nezinguquko ze-epigenetic ngokubambisana nesondlo, ukulahleka kwesisindo kunye nokungenelela kokusebenza. Kukho ubungqina obwandayo ukusuka kwizifundo zomntu kunye nezilwanyana ukuba ubudlelwane phakathi kokungcoliswa kwesondlo kunye nesifo sengqondo sokunyameka kwaye i-T2DM inokudityaniswa ngeenguqu ze-epigenetic kwinzala. Injongo yale ngongoma kukushwankathela uphuhliso olutshanje kule nsimi ehamba ngokukhawuleza, ngokugxila ngokukodwa kwi-EWAS yabantu kunye nokuphanda uphando kwimpembelelo yokutya kunye nokuphila (kokubili nangaphambi kokuzalwa) kwi-epigenome kunye nolwalamano lwawo kunye ne-metabolic iziphumo zempilo. Ubunzima bokuhlukanisa iziphumo ezivela kwiimeko ezi zifundo kunye nenxaxheba ebalulekileyo yezilwanyana zezilwanyana zokuvavanya ubudlelwane be-causal kunye nokunika ingqiqo kwiindlela ezisetyenziswayo kwakhona. Isishwankathelo, indawo ye-epigenetics kunye nempilo ye-metabolic ibone ukuphuthuma ngokukhawuleza kwithuba elifutshane. Nangona iziphumo eziza kumhla zithembisayo, uphando luyaqhubeka, kwaye ilishumi elizayo lithembisa ukuba lixesha lophando oluvelisayo kwiintsebenziswano eziyinkimbinkimbi phakathi kwe-genome, i-epigenome, kunye nommandla njengoko zihambelana nezifo ezixubileyo.
Internet: I-Epigenetics, i-DNA methylation, i-Obesity, i-2 yeswekile, iProgramming Development
Contents
Kuluntu apho ukutya okuxineneyo amandla kuninzi kwaye nesidingo sokwenza umthambo sisezantsi, kukho umahluko obanzi kubantu ngabanyeāukuba sengozini yokuphuhla ļæ½ukutyeba kakhulu kunye neengxaki zempilo ye-metabolic. Uqikelelo lwendima yofuzo kolu lwahluko lukuluhlu lwe-40ļæ½70 %, kwaye ngelixa izifundo ezinkulu ze-genome-wide association (GWAS) zichonge inani le-loci yofuzo ehambelana nomngcipheko wokutyeba, i- ~ 100 eyona nto ixhaphakileyo yokwahluka kwemfuza kuphela. akhawunti yeepesenti ezimbalwa zokungafani kokutyeba [2, 3]. Uqikelelo olubanzi lwe-genome luphezulu, lubalelwa kwi ~20 % yokwahluka [3]; nangona kunjalo, inxalenye enkulu yelifa lihlala lingachazwanga.
Kutshanje, ingqwalasela iguqukele ekuphandeni indima yotshintsho lwe-epigenetic kwi-etiology yokutyeba. Kuxoxiwe ukuba i-epigenome inokumela ikhonkco lomatshini phakathi kwezinto ezahlukeneyo zofuzo kunye nezinto ezisingqongileyo ekumiseleni umngcipheko wokutyeba kwaye inokunceda ukucacisa ļæ½ilifa elilahlekileyo.ļæ½ Izifundo zokuqala ze-epigenetic zabantu zazincinci kwaye ziphande kuphela inani elilinganiselweyo le-loci. Ngelixa oku ngokubanzi kubangele ukuphindaphinda okulambathayo, ezinye zezi ziphumo zakwangoko, umzekelo ubudlelwane phakathi kwePGC1A methylation kunye nohlobo lwe-2 yeswekile mellitus (T2DM) [4] kunye nezinye njengoko kuxoxwe ngazo kuvan Dijk et al. [5], ziye zaphindwa kwizifundo zamva. Inkqubela phambili yamva nje kunye nokwanda kokufikeleleka kwetekhnoloji ephezulu ngoku ivumela izifundo ezinkulu ze-epigenome wide association (EWAS) kunye nokudityaniswa kweengqimba ezahlukeneyo zolwazi lwe-genomic ukuhlola unxibelelwano oluntsokothileyo phakathi kwe-genotype, i-epigenome, i-transcriptome, kunye nokusingqongileyo [6] 9]. Olu phononongo lusesebuncinaneni balo, kodwa iziphumo ukuza kuthi ga ngoku zibonise isithembiso ekuncedeni ukucacisa umahluko wokutyeba kakhulu.
Kukho ubungqina obandayo bokuba ukutyeba kakhulu kuye kwavela kwimvelaphi yengqondo, njengoko ukubhengeza ukutya okunesondlo ngaphambi kokuzalwa okanye kwasebusaneni kunxulunyaniswa nokwanda komngcipheko wokutyeba kakhulu kunye nesifo se-metabolic kwimpilo kamva [10-13]. Ekuqaleni, uphononongo lwezilwanyana lubonakalise ukuba uluhlu lokuvezwa kokutya okunesondlo ebomini, ngakumbi abo banamava kwangoko ekukhulelweni, kunokubangela utshintsho lwe-epigenetic kwizicubu eziphambili zemetabolism zenzala eyaqhubeka emva kokuzalwa kwaye ikhokelela kutshintsho olusisigxina ekusebenzeni kwemfuza [13-17]. Ubungqina buyavela ukuxhasa ubukho bendlela efanayo ebantwini. Oku kukhokelele kukhangelo lweempawu ze-epigenetic ezikhoyo kwangethuba ebomini ezixela kwangaphambili umngcipheko wesifo se-metabolic, kunye nezifundo zokufumanisa ukuba ngaba inkqubo ye-epigenetic yesifo se-metabolic inokuthintelwa okanye ibuyiselwe kubomi kamva.
Olu hlalutyo lunikeza ukuhlaziywa kwenkqubo yethu yangaphambili yokuphononongwa kwezifundo kwi-epigenetics kunye nokukhuluphala kubantu [5]. Ukuphononongwa kwethu kwangaphambili kubonakalise iziphumo ezithembisayo zezifundo zokuqala, kubandakanywa amanqaku okuqala okuba nokukhuluphala okunokufunyanwa ekuzalweni (umzekelo, i-RXRA) [18]. Nangona kunjalo, kwagqitywa kwakhona ukuzaliswa kobuninzi bezinto ezifunyenweyo kunye nokungabikho kophando olude lwexesha elide. Ukuhlaziywa kwangoku kugxininise ukuphuhliswa kwangoko kwintsimi ehamba ngokukhawuleza kwaye, ngokukodwa, kwi-EWAS yabantu kunye nokuphanda uphando malunga nempembelelo ye-epigenome kunye nendima ephumayo ye-epigenetics kwi-pathology of fatness . Siphinde sibhekane nobunzima ekuboneni iimeko ezikhoyo kwezi zifundo kunye nokubaluleka kweempawu zezilwanyana ekuboneleleni ngeendlela.
Izifundo zezilwanyana zidlale indima ebaluleke kakhulu ekuqondeni kwethu ngoku kwendima ye-epigenetics kwimvelaphi yophuhliso lokutyeba kakhulu kunye ne-T2DM. Zombini ukwanda kunye nokwehla kwesondlo sikamama ngexesha lokukhulelwa kunxulunyaniswa nokunyuka kwamafutha kwinzala yezona ntlobo zininzi zezilwanyana ezanyisayo ezifundwe ukuza kuthi ga ngoku (zihlaziyiwe kwi [11, 13-15, 19]). Ukutya komama ngexesha lokukhulelwa akubanga neziphumo ezichanekileyo kwimveku engekazalwa, inokuba nefuthe ngokuthe ngqo kwii-oocyte ezikhulayo zabasetyhini ababhinqileyo kunye neeseli zentsholongwane zokuqala zesisu esiswini kwaye ke oko kunokuba nefuthe kwisiphelo kunye nenzala enkulu. Yiyo loo nto idatha yedatha eyahlukeneyo ihlala ifuneka ukwahlulahlula phakathi koomama phakathi kokuzalwa kunye neendlela zokudlulisela ezizukulwaneni.
Itheyibhile 1 isishwankathela iintlobo ezahlukeneyo zezilwanyana ezisetyenzisiweyo ukubonelela ngobungqina beenguqu zokuguqulwa kwamagciwane kunye ne-epigenetic kwinzala ehambelana nesicwangciso somzali wesondlo. Iqulethe iinkcukacha eziphathelele kwizifundo ezichonga amanqaku e-epigenetic ashintshiweyo kubantu abadala abadala abajongene nemingeni yokutya. Itafile ihlelwe luhlobo lokudlulisela umngcipheko olungcetywayo.
Ukubonisa ubungqina obuvela kwizifundo zezilwanyana kunye nokufumaneka kwezinto ezifikelelekayo zokuhlalutya kwe-genome, kuye kwandiswa ngokukhawuleza kwezifundo ze-epigenome kubantu. Ezi zifundo zijolise kakhulu ekuchongwa kweendawo ezahlukileyo kwi-DNA methylation ezidibaniswa ne-phenotypes.
Umbuzo osisiseko ngumlinganiselo apho ukuguqulwa kwe-epigenetic kunceda ekuphuhliseni i-phƩnotype ye-metabolic, kunokuba nje kulandelelanise (umzobo 1). Iiprogramme ze-epigenetic zinokubangela inxaxheba ekuphuhliseni ukunyanya, kunye nokudlala indima kwimingcipheko yengozi ye-cardiovascular and metabolic. Kwizifundo zabantu, kunzima ukubonisa ubungqina bendalo [44], kodwa ukungqinelana kungenziwa kwimiba yobungqina:
(ii) Isikhathi seenguqu ze-epigenetic. Ubukho beempawu ze-epigenetic ngaphambi kokuphuhliswa kwe-phenotype luphawu olubalulekileyo oluhambelana nobunzima. Ngakolunye uhlangothi, ubukho bamanqaku ngokubambisana nokukhuluphala, kodwa kungekhona ngaphambi kokuphuhliswa kwayo, kunokusetyenziswa ukukhupha ngaphandle kwexesha kodwa akuyi kubandakanywa indima enokuthi inxaxheba kwimeko yokugula enxulumene nokugqithisa.
(iii) Ukwahlula ngokucacileyo kwendlela. Oku kubhekisela kumatshintsho e-epigenetic anxulumene nokutshintshwa kweentlobo zegesi kunye nenxaxheba ebalulekileyo ekulawuleni i-phenotype yomdla. Omnye umzekelo onjalo ngumbutho we-methylation kwiindawo ezimbini zeCpG kwi-genetic CPT1A kunye namazinga e-triglyceride ejikelezayo [50]. I-CPT1A ifake i-carnitine palmitoyltransferase i-1A, i-enzyme eneendima ephambili kwi-fatty metabolism, ibe le nto ibonisa ukuba ukuhlukana kwemithamo ye-methylation yalo mfuzo inokuthi ihambelane neenguqu kwi-plasma triglyceride concentrations.
Uninzi lophando lwakutsha nje lugxile ekuphononongeni unxulumano phakathi kokutyeba kakhulu/izifo ze-metabolic-kunye ne-DNA methylation kuyo yonke i-genome (iThebhile 2). I-EWAS enkulu epapashiweyo ukuza kuthi ga ngoku, kubandakanywa inani labantu be-5465, ichonge iindawo ze-37 ze-methylation egazini eziye zadibaniswa ne-index mass body (BMI), kubandakanywa neziza kwi-CPT1A, ABCG1, kunye ne-SREBF1 [51]. Olunye uphononongo olukhulu lubonise imibutho engaguqukiyo phakathi kwe-BMI kunye ne-methylation kwi-HIF3A kwigazi elipheleleyo kunye nezicubu ze-adipose [52], ukufumanisa okuye kwaphinda kwaphindwa ngokuyinxenye kwezinye izifundo [9, 51]. Olunye unxulumano olusandulā ukuxelwa phakathi kwemilinganiselo enxulumene nokutyeba kakhulu kunye ne-DNA methylation ibandakanya (i) i-DNA methylation umahluko phakathi kokubhitya kunye ogqithiseleyo abantu abakwi-LY86 kwii-leukocyte zegazi [53]; (ii) imibutho phakathi kwe-PGC1A yokukhuthaza i-methylation kwigazi elipheleleyo labantwana kunye ne-adiposity kwiminyaka emi-5 kamva [54]; (iii) imibutho phakathi kwe-hip-hip ratio kunye ne-ADRB3 methylation egazini [55]; kunye (iv) nemibutho phakathi kwe-BMI, amanyathelo okuhambisa amanqatha, kunye neendawo ezininzi ze-DNA methylation kwizicubu ze-adipose [9, 56]. I-EWAS ikwabonakalise unxibelelwano phakathi kweendawo ze-DNA methylation kunye nee-lipids zegazi [55, 57-59], i-serum metabolites [60], ukumelana ne-insulin [9, 61], kunye ne-T2DM [48, 62, 63] (2 Table).
Olunye uphononongo lwamva nje lubonelele ubungqina bokuba ukutya okondlayo kokubeleka ngaphambi kokuzalwa kunye nokutyeba kakhulu okanye imeko yesifo seswekile kunxulunyaniswa notshintsho lwe-DNA methylation kwizakhi zofuzo ezinxulumene nokukhula kombungu, ukukhula, kunye nesifo se-metabolic kwimbewu [70-73].
Nangona iinkcukacha zoluntu zingqongqo, kukho izibonakaliso zokuba ukukhuluphala komntwana kungakhokelela ekuveliseni i-methylation yeengqungquthela zentsholongwane kwi-74], isiphumo esicatshangelwa ukuba sixolelane ngeenguqu ze-epigenetic ezenziwa ngexesha le-spermatogenesis.
Ngokubambisana, ubungqina bokuxhasa amandla okulungisa i-epigenome kubantu abadala bacebisa ukuba kunokubakho ithuba lokungenelela kwimpilo emva kokubeletha ukulungisa okanye ukuguqula iprogram ye-epigenetic.
Ukubaluleka kwezinto eziphilayo zenguqu ezincinci zeemethylation kuye kwacelwa. Nangona kunjalo, kwiishubhu ezenziwe ngumxube weentlobo zeeseli, utshintsho oluncinci kwi-DNA methylation lungabonakalisa utshintsho olukhulu kwiqhekeza elithile leseli. Ukuhlanganiswa kweenkcukacha ze-epigenome kunye ne-transcriptome kunye nezinye i-epigenetic idatha, ezifana nokuguqulwa kwe-histone, kubalulekile, ekubeni utshintsho lwe-DNA lwe-methylation lungabonakalisa utshintsho olukhulu kwi-chromatin structure kwaye lunokudityaniswa neenguqu ezibanzi kwimbonakalo yemfuza. Umongo we-genomic kufuneka uqwalaselwe; utshintsho oluncinci ngaphakathi kwento elawulayo efana nomgqugquzeli, umgqugquzeli, okanye umshini wokufaka isilisi unokubaluleka kokusebenza. Kule nkalo, i-DMR yokunyanyisa, kunye nemimandla echaphazelekayo yenkomfa yokubeletha kunye ne-meQTL ye-loci ye-metamic traits i-loci ziye zabonwa ukuba zithintele izinto eziphuculisayo [8, 43, 68]. Kukho ubungqina bokuba i-DNA methylation kwimimandla ejongene nendlala inokuyichaphazela into eyenziwa yintsebenziswano [68], exhasa indima yesondlo-i-methyl
Umda omkhulu kwizifundo ezininzi zabantu kukuba iimpawu ze-epigenetic zivame ukuvavanywa kwigazi le-peripheral, kunokuba kwizicubu ezifanelekileyo ze-metabolically (Umfanekiso 2). I-heterogeneity yegazi ngumba, kuba iiseli ezahlukeneyo zineesignitsha ezahlukeneyo ze-epigenetic, kodwa i-algorithms iphuhlisiwe ukuqikelela ukubunjwa kweeselula ukunqoba le ngxaki [86]. Mhlawumbi okona kubaluleke kakhulu, amanqaku e-epigenetic kwiiseli zegazi asenokungaxeleli imeko yezicubu zomdla ophambili. Nangona kunjalo, uphando olutshanje lunikeze ubungqina obucacileyo bobudlelwane phakathi kwamanqaku e-epigenetic kwiiseli zegazi kunye ne-BMI. Kwimeko ye-HIF3A apho inqanaba le-methylation (i-beta-value) kubantu abafundayo ukusuka kwi-0.14ļæ½0.52, i-10% yokunyuka kwe-methylation idibene nokunyuka kwe-BMI ye-7.8% ļæ½[52]. Ngokufanayo, umahluko we-10% kwi-PGC1A methylation unokuqikelela ukuya kuthi ga kwi-12% umahluko kubunzima bamafutha [54].
Ukufundwa kwendima ye-epigenetics ekugqithiseni nokugqithisa izifo kwandiswe ngokukhawuleza kwiminyaka yamuva nje, kwaye ubungqina bubuthelela ukudibanisa phakathi kokuguqulwa kwe-epigenetic kunye neziphumo zempilo ze-metabolic kubantu. Iingxaki ze-epigenetic biomarker ezinxulumene nokukhuluphala kunye nempilo yokuxilonga ziye zavela kwizifundo zamuva. Ukuqinisekiswa kwamagqabha epigenetic kumaqela amaninzi, ukuba amanqaku amaninzi atholakala kwiijethi ezinomsebenzi onobunzima bokuphuhliswa nokuphuhliswa kwe-T2DM, kunye nokugqithwa kwamanqaku epigenetic kunye nokunyameka okwaziwayo kunye ne-T2DM lotic genetic kuqinisa ubungqina bokuba le mibutho ngokwenene. Ubunzima kubunzima ukuseka; Nangona kunjalo, kungakhathaliseki ukuba iminyano ibangela, amanqaku e-epigenetic achongiweyo anokuthi afanele afaneleke njengama-biomarkers ukunyanya kunye nomngcipheko wesifo somzimba.
Iziphumo zobukhulu kwizicwili ezifikeleleka lula ezifana negazi zincinci kodwa zibonakala ngathi ziyaveliswa ngaphandle kokwahluka kubuzwe, uhlobo lwethishu, kunye neendlela zohlalutyo [51]. Kwanaxa utshintsho lwe-methylation encinci ye-DNA inokubaluleka kwendalo. Indlela yokudibanisa i- omics iya kubaluleka ekuchazeni ngakumbi ukusebenzisana okunzima phakathi kwe-epigenome, umbhalo okhutshelweyo, ufuzo, kunye nempilo ye-metabolic. Izifundo ezide, ezichaphazela izizukulwana ezininzi, zibalulekile ekumiseni ubudlelwane obunobangela. Singalindela ezinye izifundo ezinjalo kwixa elizayo, kodwa oku kuya kuthatha ixesha.
Nangona uphando lwezilwanyana luqhubeka lubonisa umphumo wobomi bokuqala zesondlo Ukuvezwa kwi-epigenome kunye nempilo ye-metabolic yenzala, idatha yomntu isenomda. Nangona kunjalo, uphando olutshanje lunikeze ubungqina obucacileyo bokuthi ukuvezwa kwisondlo esincinci ngexesha elithile lokuphuhliswa kwangaphambi kokubeletha kuhambelana notshintsho lwe-methylation kwinzala kwaye ngoko ke unakho ukuchaphazela i-phenotype yabantu abadala. Izifundo zezilwanyana ziya kubaluleka ukuqinisekisa ukufunyaniswa kwabantu kwindawo elawulwa ngakumbi, kunceda ukufumanisa ukuba utshintsho oluchongiweyo lwe-methylation lunempembelelo kwimpilo ye-metabolic, kunye nokutyhila iindlela eziphantsi komgaqo we-epigenetic phakathi kwezizukulwana / ze-transgenerational. Ukuchongwa kweendlela ze-causal eziphantsi kweempendulo zememori ye-metabolic, indlela yokudluliselwa kweziphumo ze-phenotypic kwizizukulwana ezilandelelanayo, iqondo lempembelelo kunye nokuzinza kweempawu ezidluliswayo, kunye nokuchongwa komxholo obanzi kunye nokumanyanisayo wendaleko kukwahlala kuyimibuzo ebalulekileyo ekufuneka iqwalaselwe. . Le yokugqibela isoloko igqunyelelwe yingqikelelo yempendulo eguqukayo exelwe kwangaphambili, okt, impendulo kwimeko-bume elindelweyo yexesha elizayo eyandisa ukomelela kwabantu. Nangona kunjalo, le ngcamango iye yathandabuza kakhulu njengoko kukho ubungqina obulinganiselweyo bokukhula kokuqina komzimba kamva ebomini [87].
Ngesishwankathelo, iziphumo zithembisa, njengoko utshintsho lwe-epigenetic ludibaniswa nempilo yabantu abadala kunye nokusebenza njengomlamli phakathi kokutya kokubeleka kokubeleka kunye nokwanda komngcipheko weziphumo zempilo ezingekho mzimba. Amatshwa epigenetic amatsha afunyenwe adibene namanyathelo empilo yokuxilonga. Ukuhlanganiswa kwezigaba ezahlukeneyo zolwazi lwe-genomic kuye kwandise inkxaso engakumbi kwiimeko zokubambisana, kwaye kukho izifundo ezongezelelekileyo ezibonisa iziphumo zendalo yangaphambili kunye neyokuzalwa emva kwe-epigenome kunye nempilo. Nangona kukho imibuzo ebalulekileyo ehleliyo, ukuqhubela phambili kwindlela yokwenziwa kwezinto ezikhoyo kwangoku kwenze ukuba iindidi zezifundo ezininzi ezijoliswe ngabantu ziza kufuneka ukuba zijongane nezikhewu zolwazi. Iiminyaka elishumi ezayo ithembisa ukuba yithuba lomsebenzi omkhulu kule ndawo ebalulekileyo yophando.
USusan J. van Dijk1, uRoss L. Tellam2, uJanna L. Morrison3, uBeverly S. Muhlhausler4,5 noPeter L. Molloy1 *
Ukunyanzela umdla
Ababhali bavakalisa ukuba abanalo inxaxheba.
Igalelo lababhali
Bonke ababhali babenegalelo ekuqulunqweni nasekuhlaziyweni okubalulekileyo kwincwadi yesandla, kwaye bonke abalobi bafunde baze bavunywa umbhalo wesigqi wokugqibela.
Ulwazi lwababhali
UBeverly S. Muhlhausler kunye noPeter L. Molloy babambene nabalobi bokugqibela.
Imibulelo
Lo msebenzi usexhaswe yigranti evela kwiNgxowa-mali yeSayensi kunye neNkampani (Grant RP03-064). I-JLM kunye ne-BSM zixhaswa yiNational Health and Medical Research Council (CBL), i-APP1066916; i-BSM, i-APP1004211). Siyabulela uLance Macaulay noSue Mitchell ukuze bafunde ngokubalulekayo kunye namazwana kwi-text.
Iinkcukacha zobhali
I-1CSIRO Ukutya kunye neZondlo zeFlagship, i-PO Box 52, iNorth Ryde, i-NSW 1670, i-Australia. I-2CSIRO I-Flagship yezoLimo, i-306 ye-Carmody Road, i-St Lucia, i-QLD 4067, i-Australia. I-3Imvelaphi yokuqala yeQela loPhando lwezeMpilo yaBadala, iSikolo seKhemesti kunye neSayensi yezoNyango, iSansom Institute yoPhando lwezeMpilo, iYunivesithi yaseMzantsi Australia, i-GPO Box 2471, i-Adelaide, i-SA 5001, i-Australia-4FOODplus Research Centre, i-Waite Campus, iYunivesithi yase-Adelaide, i-PMB 1, Glen Osmond, SA 5064, Australia. 5Iziko loPhando ngeMpilo yabasetyhini kunye nabantwana, 72 King William Road, North Adelaide, SA 5006, Australia.
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mngcipheko_abacwangcisi / ukwephuza ngokweqile / en / index.html. Ifumaneka 29 Januwari 2015.
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Ubume obubambeneyo: ILayisensi yeeNkcazo ezininzi: Ugunyaziswe Ukuziqhelanisa I-40 States*
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