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Start arrow Autosomalne recesywne arrow Metabolizm Fenyloalaniny

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Metabolizm Fenyloalaniny Utwórz PDF Drukuj Wyślij znajomemu
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Fenyloalanina dla człowieka jest aminokwasem egzogennym, a jej główne źródła stanowią białka pokarmowe. W warunkach fizjologicznych 75 % wchłoniętej do krwiobiegu fenyloalaniny ulega przemianie (hydroksylacji) do aminokwasu tyrozyny w komórkach wątroby
Fenyloalanina dla człowieka jest aminokwasem egzogennym, a jej główne źródła stanowią białka pokarmowe. W warunkach fizjologicznych 75 % wchłoniętej do krwiobiegu fenyloalaniny ulega przemianie (hydroksylacji) do aminokwasu tyrozyny w komórkach wątroby.
Jest to główny szlak przemiany fenyloalaniny w ustroju, który zachodzi w warunkach tlenowych pod obecność wątrobowego enzymu hydroksylazy fenyloalaniny oraz kofaktora tej reakcji - tetrahydrobiopteryny (BH4). Blok przemiany fenyloalaniny do tyrozyny w przebiegu fenyloketonurii jest spowodowany całkowitym lub częściowym niedoborem PAH (Rys. 1).
 
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Zaburzenie hydroksylacji fenyloalaniny prowadzi do nagromadzenia jej w dużych ilościach we krwi, w płynach ustrojowych oraz ośrodkowym układzie nerwowym (o.u.n.). Nagromadzenie olbrzymich ilości fenyloalaniny w płynie mózgowo-rdzeniowym i tkance nerwowej stanowi podstawową przyczynę uszkodzenia układu nerwowego u chorych nie leczonych, a przejawia się głównie upośledzeniem umysłowym.


Badania mechanizmów oddziaływania wysokich poziomów fenyloalaniny na ośrodkowy układ nerwowy (o.u.n.) stały się możliwe z chwilą stworzenia modelu niedoboru PAH u myszy laboratoryjnych PAH

Patomorfologia i Patofizjologia upośledzenia umysłowego
Badania mechanizmów oddziaływania wysokich poziomów fenyloalaniny na ośrodkowy układ nerwowy (o.u.n.) stały się możliwe z chwilą stworzenia modelu niedoboru PAH u myszy laboratoryjnych PAH.
Jak są przekazywane mutacje?

Fenyloketonuria jest chorobą genetyczną, dziedziczoną od rodziców autosomalnie recesywnie. Oznacza to, że każdy z rodziców na jednym z chromosomów autosomalnych 12-tej pary posiada jedną kopię zmutowanego genu hydroksylazy fenyloalaniny (PAH) - "nieaktywny gen"- i jest nosicielem danej cechy (niedoboru lub braku hydroksylazy fenyloalaniny).
Nie prowadzi to do ujawnienia się choroby (cechy) u nosicieli, ale mogą oni przekazywać cechę tę następnym pokoleniom (swoim dzieciom). Jeżeli potomstwo odziedziczy tylko jeden "nieaktywny gen"  - staje się nosicielem, u niego również nie będą występować objawy niedoboru PAH.
W przypadku gdy dziecko odziedziczy od każdego rodzica po jednej kopii chromosomu 12-tego ze zmutowanym genem PAH, dochodzi do ujawnienia się bloku enzymatycznego niedoboru hydroksylazy fenyloalaniny.

W wyniku przeprowadzonych prac doświadczalnych oraz obserwacji wysnuto hipotezy, pozwalające tłumaczyć mechanizm powstawania upośledzenia umysłowego u osób z PKU, które nie były leczone, były leczone niedostatecznie lub leczenie zakończono z różnych powodów.
W przebiegu hiperfenyloalaninemii spowodowanej niedoborem PAH brane są pod uwagę następujące prawdopodobne mechanizmy strukturalnego oraz  czynnościowego uszkodzenia o.u.n. (bariery krew-mózg):
"    zaburzenia transportu dużych aminokwasów przez barierę krew-mózg w przebiegu hyperfenyloalaninemii we krwi obwodowej
"    obniżenie produkcji neurotransmitterów
"    zaburzenia syntezy i metabolizmu mieliny
Jak są przekazywane mutacje?

Fenyloketonuria jest chorobą genetyczną, dziedziczoną od rodziców autosomalnie recesywnie. Oznacza to, że każdy z rodziców na jednym z chromosomów autosomalnych 12-tej pary posiada jedną kopię zmutowanego genu hydroksylazy fenyloalaniny (PAH) - "nieaktywny gen"- i jest nosicielem danej cechy (niedoboru lub braku hydroksylazy fenyloalaniny).

Od genu do enzymu


W wyniku odczytu prawidłowej informacji z genu PAH i transformacji jej na polecenie syntezy białka powstaje jeden łańcuch (monomer) enzymu PAH, który zawiera 451 aminokwasów. Monomery łącząc się tworzą funkcjonalny enzym PAH. Dokładna liczba podjednostek i struktura trójwymiarowa enzymu nie są znane.
Zmiany w strukturze genu PAH (mutacje) mają znaczący wpływ na niską syntezę, defekt lub brak docelowego produktu, czyli czynnego enzymu. W wyniku mutacji genu hydroksylazy fenyloalaniny ulega zmianie jego struktura, co prowadzi do utraty lub obniżenia jego aktywności. Zmieniony strukturalnie oraz o niskiej aktywności enzym nie spełnia swojej roli w procesie hydroksylacji fenyloalaniny.
 
Mutacje w genie PAH są przyczyną niedoboru PAH i reprezentują stałą jednostkę chorobową z obrazem klinicznym hiperfenyloalaninemii we krwi obwodowej i ośrodkowym układzie nerwowym. Większość z tych mutacji jest ściśle związana z określonymi populacjami. Badania DNA pacjentów z niedoborem PAH umożliwiaj wgląd w molekularne mechanizmy będące podłożem fenotypu hyperfenylalaninemii.
 


Mutacje nowe czy stare?


Od chwili zidentyfikowania pierwszej mutacji w genie PAH baza danych zawierająca informacje dotyczące mutacji w tym genie znacznie się poszerzyła. Obecnie znanych jest ponad 420 mutacji. Występowanie ograniczonej liczby najczęściej spotykanych mutacji w większości populacji sugeruje, że gen PAH niełatwo ulega modyfikacjom oraz że są one przekazywane z pokolenia na pokolenie w niezmienionej postaci.
Z przeprowadzonych przez Flemming'a i Guttler'a w Kopenhadze prac badawczych wynika że np. przodkowie pacjentów z mutacją IVS 12ntl zamieszkiwali okolice położone na zachodnim wybrzeżu Jutlandii. Z wiosek tych ruszały liczne wyprawy duńskich wikingów do wybrzeży Anglii i północno-zachodniej Europy pozostawiając po sobie informację genetyczną.
 
Historycznie i archeologicznie zostało też potwierdzone, że rozkład allela R408W/Htl w Szkocji, Anglii i Francji ściśle jest powiązany z wyprawami i osiedleniem się irlandczyków na tych terenach między III a IV wiekiem n.e. Względnie wysoka częstość występowania allela R408W/Ht2 wśród genetycznie zróżnicowanych narodów słowiańskich, bałtyckich i germańskich sugeruje ekspansję tej mutacji poprzez dyfuzję (przenikanie) demograficzną.
 
Najważniejszą więc rolę w rozpowszechnianiu mutacji PAH odgrywała migracja ludzi a nie selekcja, dlatego zupełnie odmienne mutacje występują obecnie w północnych (R413P) i w południowych (IVS 4ntl) Chinach, rozdzielonych rzeką Yangtze, która stanowiła olbrzymią przeszkodę migracyjną w dawnych czasach.
 
Przytoczone wyżej przykłady są małą częścią prowadzonych badań genealogicznych, historycznych i archeologicznych, na podstawie których opracowano szereg map ilustrujących drogi "migracji" mutacji w genie PAH w skali światowej. Powstały również fizyczne mapy genu PAH lokalizujące każdą z dotychczas znanych mutacji.
 
Czy wszystkie mutacje w genie PAH powodują występowanie PKU u ludzi?


Nie wszystkie mutacje w genie PAH powodują występowanie PKU, część spośród znanych mutacji jest bezobjawowa ("silent"). Pozostałe mutacje powodują podstawowe dla PKU zaburzenie - niedobór enzymu PAH, który prowadzi do wzrostu stężenia fenyloalaniny w surowicy krwi.
Wartości oznaczanych stężeń fenyloalaniny w surowicy krwi u noworodków wyłonionych w przesiewie noworodkowym w kierunku PKU wahają się w szerokich granicach od 6 do > 40 mg/dl (360 do > 2400 µmol/l). Jest to spowodowane różnorodnością dziedziczonych przez dzieci mutacji w genie PAH, które w zróżnicowany sposób ograniczają aktywność hydroksylazy fenyloalaniny. Wyróżnia się trzy grupy mutacji: silne (S), pośrednie (P) i łagodne (Ł).
Znając rodzaj mutacji w genie PAH (genotyp - podłoże genetyczne) można przewidywać ciężkość przebiegu klinicznego fenyloketonurii, czyli jej obraz metaboliczny (fenotyp - przełożenie funkcji genu na obraz chorobowy), inaczej mówiąc korelację genotyp-fenotyp.
 
W przypadku występowania u dziecka silnych mutacji (S+S) można się spodziewać fenotypu fenyloketonurii klasycznej - phe > 20 mg/dl (1200 µmol/l), silnej i pośredniej (S+P) oraz pośrednich (P+P) oczekujemy fenyloketonurii łagodnej - phe 10-20 mg/dl (600-1200 µmol/l) rzadko łagodnej hyperfenyloalninemii, innych wariantów (S+Ł), (P+Ł), (Ł+Ł) przewidujemy łagodną hyperfenyloalaninemię - phe 2-10 mg/dl (120-600 µmol/l) lub fenotyp nieodróżnialny od prawidłowego.
 

Podwyższone stężenie fenyloalaniny - postępowanie terapeutyczne



Po stwierdzeniu u dziecka podwyższonego stężenia fenyloalaniny (hiperfenyloalaninemii) w teście bibułowym (kolorymetrycznym), pobranym w 3 dobie życiapo pełnym obciążeniu dziecka białkiem pokarmowym, rodzice/opiekunowie dziecka otrzymują pisemne wezwanie do ośrodka referencyjnego. W trakcie pierwszej wizyty w specjalistycznej poradni rodzicom/opiekunom dziecka udzielana jest informacja odnośnie istoty podejrzewanej choroby, przyczyn jej występowania, możliwości leczenia.
Wobec istnienia wielu przyczyn hiperfenyloalaniemii: fenyloketonuria, nietypowe postacie fenyloketonurii oraz tyrozynemia, niezbędne jest ich różnicowanie. Taka diagnostyka różnicowa oraz ewentualne wdrożenie leczenia odbywają się w warunkach szpitalnych. W oparciu o wyniki testów diagnostycznych, wykonanych natychmiast po przyjęciu dziecka do oddziału, podejmowana jest decyzja o dalszym leczeniu. W przypadku potwierdzenia niedoboru hydroksylazy fenyloalaniny (fenyloketonuria) wdrażane jest leczenie dietetyczne. Natomiast wobec stwierdzenia innego rodzaju defektu metabolizmu obierane są inne metody lecznicze.
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