Ó÷åáíîå ïîñîáèå: Îáì³í âóãëåâîä³â
Ó÷åáíîå ïîñîáèå: Îáì³í âóãëåâîä³â
Ïëàí
1. Ôåðìåíòàòèâíå ïåðåòâîðåííÿ âóãëåâîä³â ó õàð÷îâîìó òðàêò
ëþäèíè
2. Ïîøèðåííÿ âóãëåâîä³â ó îðãàí³çì³ ëþäèíè
3. Ïðîöåñè ïåðåòâîðåííÿ âóãëåâîä³â íà êë³òèííîìó ð³âí³:
3.1. Ñèíòåç ãë³êîãåíó
ç ãëþêîçè (ãë³êîãåíåç)
3.2. Ðîçïàä ãë³êîãåíó äî ãëþêîçè (ãë³êîãåíîë³ç)
3.3. Ãë³êîë³ç (àíàåðîáíèé ðîçïàä)
3.4. Äèõàííÿ (àåðîáíèé ðîçïàä âóãëåâîä³â)
4. Åíåðãåòè÷í³ åôåêòè ïðîöåñ³â
Âóãëåâîäè ñêëàäàþòü á³ëÿ 70 % õàð÷îâîãî ðàö³îíó ëþäèíè (áîðîøíî,
êðóïè, êàðòîïëÿ – êðîõìàëåâì³ñí³; ìåä, ÿãîäè, ôðóêòè – ì³ñòÿòü ñàõàðîçó, ãëþêîçó
ôðóêòîçó), ÿâëÿþ÷èñü îñíîâíèì äæåðåëîì åíåð㳿 äëÿ îðãàí³çìó. Ïîøèðåíèé ó
ïðèðîä³ ïîë³ñàõàðèä êë³òêîâèíà ò³ëüêè ÷àñòêîâî çàñâîþºòüñÿ îðãàí³çìîì ëþäèíè çà
äîïîìîãîþ áàêòåð³é òîâñòîãî êèøå÷íèêà. Îñíîâíîþ ôóíêö³ºþ êë³òêîâèíè
çàáåçïå÷åííÿ ïåðèñòàëüòèêè õàð÷îâîãî òðàêòó, ñòèìóëþâàííÿ ñåêðåòîðíî
ä³ÿëüíîñò³, ñïðèÿííÿ ñèíòåçó â³òàì³íó Ê çà äîïîìîãîþ êèøå÷íî¿ ì³êðîôëîðè.
Ïîñê³ëüêè âóãëåâîäè çàñâîþþòüñÿ ó âèãëÿä³ ìîíîöóêðèä³â, òî ñêëàäí
âóãëåâîäè (äèöóêðèäè, ïîë³öóêðèäè) ïîâèíí³ ïîïåðåäíüî ðîçùåïèòèñÿ. óäðîë³òè÷íå
ðîçùåïëåííÿ ïðîõîäèòü ï³ä âïëèâîì ôåðìåíò³â ùî íîñÿòü çàãàëüíó íàçâó ãë³êîçèäàçè.
Îïòèìóì 䳿 ãë³êîçèäàç ëåæèòü â íåéòðàëüíîìó òà ñëàáêîëóæíîìó ñåðåäîâèù
(âèêëþ÷åííÿ àì³ëàçà ñëèíè – ñëàáêîêèñëå), ùî ³ îáóìîâëþº ïåâíó ëîêàë³çàö³þ
ïðîöåñ³â ðîçùåïëåííÿ ó õàð÷îâîìó òðàêò³ ëþäèíè.
³ää³ë
òðàâëåííÿ
|
Îïòèìóì 䳿 ôåð-ìåíòíèõ
ñèñòåì (ðÍ) |
Ïðîöåñè |
Ôåðìåíòè |
Ðîòîâà ïîðîæíèíà |
6.5–7.0 |
Ðîçùåïëåííÿ êðîõìàëþ äî
äåêñòðèí³â, ÷àñòêîâî ìàëüòîçè äî
ãëþêîçè ³ ôðóêòîçè. |
àì³ëàçà ñëèíè,
ìàëüòàçà. |
Ñòðàâîõ³ä |
7 |
Ïðîöåñè ðîçùåïëåííÿ
â³äñóòí³ ó çâ‘ÿçêó ç øâèäêèì ïðîõîäæåííÿì ñóáñòðàò³â. |
|
Øëóíîê |
1–2 |
Ïðîöåñè ðîçùåïëåííÿ
â³äñóòí³ ó çâ‘ÿçêó ç ³íàêòèâàö³ºþ ôåðìåíò³â «êèñëèì» ñåðåäîâèùåì. |
|
12–ïàëà êèøêà |
7.0–7.5 |
Îñòàòî÷íå ðîçùåïëåííÿ
äåêñòðèí³â äî äèñàõàðè-ä³â. |
ôåðìåíòè ï³äøëóíêîâî
çàëîçè: ê³íöåâà äåêñòðèíàçà, –àì³ëàçà, ìàëüòàçà,
ñàõàðàçà, ëàêòà-çà.
|
Òîíêèé êèøå÷íèê |
7.0–7.5 |
Ðîçùåïëåííÿ äèñàõàðèä³â
äî ìîíîñàõàðèä³â. Ïîñòó-ïàííÿ ìîíîñàõàðèä³â ó êðîâ. |
ʳíöåâà äåêñòðèíàçà, –àì³ëàçà, ìàëüòàçà,
ñàõàðàçà, ëàêòà-çà.
|
Òîâñòèé êèøå÷íèê |
|
Ðîçùåïëåííÿ êë³òêîâèíè
äî öåëîá³îçè ³ ãëþêîçè. Ñèíòåç â³òàì³íó Ê. Ïðîõî-äÿòü ïðîöåñè çáðîäæåííÿ
âóãëåâîäíåâèõ çàëèøê³â ç âèä³ëåííÿì ÑÎ2, Í2, ÑÍ4.
Ôîðìóâàííÿ êàëîâèõ ìàñ.
|
Öåëþëàçà, ôåðìåíòí
ñèñòåìè áàêòåð³é. |
 òîíêîìó êèøå÷íèêó â³äáóâàþòüñÿ ïðîöåñè âñìîêòóâàííÿ ìîíîñàõàðèä³â ÷åðåç
ñèñòåìó ìåìáðàí ó êðîâ‘ÿíå ðóñëî. Äëÿ ïðîöåñó âñìîêòóâàííÿ õàðàêòåðíî: çíà÷-í³ åíåðãåòè÷í³ âèòðàòè îðãàí³çìó (àêòèâíèé ô³ç³îëîã³÷íèé
ïðîöåñ); ³çîìåðèçàö³ÿ (òðàíñôîðìàö³ÿ, ïåðåòâîðåííÿ) ñóì³ø³ ïðîñòèõ âóãëåâîä³â
(98 %) ó ãëþêîçó; òðàíñïîðòóâàííÿ ãëþêîçè ÷åðåç ìåìáðàíè ó ôîðì
ãëþêîçî-ôîñôàòó (íàâ³äü ïðîòè çíà÷åíü ¿¿ êîíöåíòðàö³¿ ó êðîâ‘ÿíîìó ðóñë³).
Ç òîêîì êðîâ³ ìîíîöóêðèäè ÷åðåç âîðîòíó âåíó ïîòðàïëÿþòü ó ïå÷³íêó, äå
âèêî-ðèñòîâóþòüñÿ äëÿ á³îñèíòåçó ãë³êîãåíó (3–5 %),
á³îñèíòåçó æèð³â (30–35 %), îêèñëþþòüñÿ äî âóãëåêèñëîãî ãàçó ³ âîäè (60–70 %).
Ïåâíà, ô³ç³îëîã³÷íî íîðìàëüíà ê³ëüê³ñòü ãëþêîçè (120 ìã íà 100 ñì3),
íåâ³ä‘ºìíîþ ñêëàäîâîþ êðîâ³, çàáåçïå÷óþ÷è ãîìåîñòàç îðãàí³çìó. Âîíà íåîáõ³äíà
äëÿ íîðìàëüíîãî ôóíêö³îíó-âàííÿ ãîëîâíîãî ìîçêó, ì‘ÿç³â
ñåðöÿ, íåðâîâî¿ òêàíèíè (ãëþêîçà –ãîëîâíå äæåðå-ëî
åíåð㳿).
Ïðè ï³äâèùåí³é êîíöåíòðàö³¿ ãëþêîçè â êðîâ³ (á³ëüøå 140 ìã) ãîâîðÿòü ïðî ã³ïåðãë³êåì³þ.
Âèêëèêàºòüñÿ âæèâàííÿì ïðîäóêò³â áàãàòèõ íà âóãëåâîäè (àë³ìåíòàðíà
ã³ïåðãë³êåì³ÿ); íåäîñòàòí³ì ñèíòåçîì ³íñóë³íó (öóêðîâèé ä³àáåò); ïîíàä-íîðìîâèì ñèíòåçîì àäðåíàë³íó ³ ãëþêàãîíó. Ñòàí ã³ïåðãë³êåì
ïðèâîäèòü äî îæèð³ííÿ.
Ïðè ïîíèæåí³é êîíöåíòðàö³¿ ãëþêîçè â êðîâ³ (ìåíøå 120 ìã) ãîâîðÿòü ïðî ã³ïîãë³êåì³þ.
Âèêëèêàºòüñÿ òðèâàëèì ãîëîäóâàííÿì; âåëèêèìè ô³çè÷íèìè íàâàíòàæåííÿìè;
íåäîñòàòí³ì ñèíòåçîì ãëþêàãîíó ³ àäðåíàë³íó; ïîíàäíîðìîâèì ñèíòåçîì ³íñóë³íó.
Ïðèâîäèòü äî äåñòðóêòèâíèõ çì³í â ãîëîâíîìó ìîçêó, íåðâîâ³é òêàíèí³, íèðêàõ.
Ñòàí òðèâàëî¿ ã³ïîãë³êå쳿 ïðèâîäèòü äî âèñíàæåííÿ îðãàí³çìó ³ ñìåðò³.
Îòæå:
Îñíîâíà ðîëü â ïðîöåñàõ îáì³íó âóãëåâîä³â íàëåæèòü ïå÷³íö³, ä³ÿëüí³ñòü
ÿêî¿ ðåãóëþºòüñÿ íåðâîâîþ ³ åíäîêðèííîþ ñèñòåìàìè (³íñóë³í,
àäðåíàë³í–ãëþêàãîí).
Ó âèïàäêó ïîâíîãî åíåðãåòè÷íîãî çàáåçïå÷åííÿ îðãàí³çìó, ç íàäëèøêó
ãëþêîçè ñèíòåçóºòüñÿ ãë³êîãåí. Ïðîöåñè ãë³êîãåíåçó ïðîõîäÿòü â êë³òèíàõ ð³çíèõ
îðãàí³â ³ òêàíèí, àëå íàéá³ëüøó çäàòí³ñòü äî íàêîïè÷åííÿ ìàþòü êë³òèíè ïå÷³íêè.
Ãë³êîãåí ïå÷³íêè º îñíîâíèì ðåçåðâíèì äæåðåëîì âóãëåâîä³â äëÿ îðãàí³çìó.
3. Ïðîöåñè ïåðåòâîðåííÿ âóãëåâîä³â íà êë³òèííîìó ð³âí³
3.1 Ãë³êîãåíåç (ñèíòåç ãë³êîãåíó ç ãëþêîçè)
1. Ãëþêîçà, ÿêà ç òîêîì êðîâ³ íàäõîäèòü ó ïå÷³íêó àêòèâóºòüñÿ
çà äîïîìîãîþ ôåðìåíòó ãåêñîê³íàçè ³ ÀÒÔ, ïåðåòâîðþþ÷èñü â ãëþêîçî-6-ôîñôàò:
![](/image/16236_2_1.png) HO OH
![](/image/16236_4_1.png) CH2OH
CH2O–P=O
![](/image/16236_5_1.png) ![](/image/16236_5_2.png) ![](/image/16236_5_3.png) ![](/image/16236_5_4.png) ![](/image/16236_5_5.png) ![](/image/16236_6_1.png) ![](/image/16236_7_1.png) ![](/image/16236_8_1.png) ![](/image/16236_8_2.png) ![](/image/16236_6_2.png) ![](/image/16236_7_2.png) ![](/image/16236_5_6.png) ![](/image/16236_5_7.png) ![](/image/16236_5_8.png) ![](/image/16236_5_9.png) ![](/image/16236_5_10.png) ![](/image/16236_6_3.png) ![](/image/16236_7_3.png) ![](/image/16236_8_3.png) ![](/image/16236_8_4.png) ![](/image/16236_6_4.png) H H
H ÀÒÔ ÀÄÔ H H
H
![](/image/16236_9_1.png)
OH OH H OH OH OH H
OH
H OH
H OH
Ãëþêîçà Ãëþêîçî–6–ôîñôàò
2. Ãëþêîçî-6-ôîñôàò ï³ä 䳺þ ôîñôîãëþêîìóòàçè ïåðåòâîðþºòüñÿ íà
ãëþêîçî-1-ôîñôàò:
![](/image/16236_12_1.png) HO OH
![](/image/16236_4_3.png) CH2O–P=O
CH2OH
![](/image/16236_5_11.png) ![](/image/16236_5_12.png) ![](/image/16236_5_13.png) ![](/image/16236_5_14.png) ![](/image/16236_5_15.png) ![](/image/16236_6_5.png) ![](/image/16236_7_5.png) ![](/image/16236_8_5.png) ![](/image/16236_8_6.png) ![](/image/16236_6_6.png) ![](/image/16236_7_6.png) ![](/image/16236_5_16.png) ![](/image/16236_5_17.png) ![](/image/16236_5_18.png) ![](/image/16236_5_19.png) ![](/image/16236_5_20.png) ![](/image/16236_6_7.png) ![](/image/16236_7_7.png) ![](/image/16236_8_7.png) ![](/image/16236_8_8.png) ![](/image/16236_6_8.png) H H
H H H H
ôîñôîãëþêîìóòàçà OH
![](/image/16236_14_1.png) OH OH
H OH OH OH H O–P=O
H OH
H OH OH
Ãëþêîçî–6–ôîñôàò Ãëþêîçî–1–ôîñôàò
3. Ãëþêîçî-1-ôîñôàò
ïîâòîðíî àêòèâóºòüñÿ ÓÒÔ (óðèäèíòðèôîñôàòîì) ç âèêî-ðèñòàííÿì åíåð㳿 äâîõ ìàêðîåðã³÷íèõ
çâ‘ÿçê³â, óòâîðþþ÷è àêòèâîâàíó óðè-äèí-äèôîñôàòãëþêîçó.
ÓÐÈÄÈÍ
![](/image/16236_15_1.png) ![](/image/16236_15_2.png) ![](/image/16236_16_1.png) ![](/image/16236_17_1.png) ![](/image/16236_4_5.png) CH2OH
CH2OH OH O
![](/image/16236_18_1.png) ![](/image/16236_5_21.png) ![](/image/16236_5_22.png) ![](/image/16236_5_23.png) ![](/image/16236_5_24.png) ![](/image/16236_5_25.png) ![](/image/16236_6_9.png) ![](/image/16236_7_9.png) ![](/image/16236_8_9.png) ![](/image/16236_8_10.png) ![](/image/16236_6_10.png) ![](/image/16236_7_10.png) ![](/image/16236_5_26.png) ![](/image/16236_5_27.png) ![](/image/16236_5_28.png) ![](/image/16236_5_29.png) ![](/image/16236_5_30.png) ![](/image/16236_6_11.png) ![](/image/16236_7_11.png) ![](/image/16236_8_11.png) ![](/image/16236_8_12.png) ![](/image/16236_6_12.png) H H
H H H H P
![](/image/16236_19_1.png) ![](/image/16236_20_1.png) OH ÓÒÔ Í4Ð2Î7 O
![](/image/16236_21_1.png) ![](/image/16236_22_1.png) ![](/image/16236_23_1.png) OH OH
H O–P=O OH OH H O–P=O
H OH OH
H OH OH
Ãëþêîçî–1–ôîñôàò ÓÄÔ–ãëþêîçà
4. Ïîñê³ëüêè ñèíòåç ãë³êîãåíó ïðîõîäèòü «çàòðàâî÷íèì» ìåòîäîì (ó ðîë
çàòðàâêè âèêîðèñòîâóºòüñÿ çàëèøîê ìîëåêóëè ãë³êîãåíó), òî àêòèâîâàíà
ÓÄÔ-ãëþêîçà ïðèºäíóºòüñÿ äî çàòðàâêè çà äîïîìîãîþ ãë³êîãåí-ñèíòåòàçíèõ (óòâîðþ
1,4–çâ‘ÿçêè) òà òðàíñãëþêîçèëàçíèõ (óòâîðþº 1,6–çâ‘ÿçêè) ôåðìåíòíèõ ñèñòåì,
çá³ëü-øóþ÷è ìîëåêóëó ãë³êîãåíó íà îäíó ìîëåêóëó ãëþêîçè:
ÓÐÈÄÈÍ
![](/image/16236_15_3.png) CH2OH OH
![](/image/16236_25_1.png) ![](/image/16236_25_2.png) ![](/image/16236_26_1.png) ![](/image/16236_5_31.png) ![](/image/16236_5_32.png) ![](/image/16236_5_33.png) ![](/image/16236_5_34.png) ![](/image/16236_5_35.png) ![](/image/16236_6_13.png) ![](/image/16236_7_13.png) ![](/image/16236_8_13.png) ![](/image/16236_8_14.png) ![](/image/16236_6_14.png) H H
H P Î
![](/image/16236_19_2.png) O (Ñ6H10O5)n ÓÄÔ
![](/image/16236_27_1.png) ![](/image/16236_21_2.png) ![](/image/16236_22_2.png) OH OH H O–P=O
(C6H10O5)n+1
H OH OH
ÓÄÔ–ãëþêîçà çàòðàâî÷íèé
ãë³êîãåí
Âèñíîâîê:
ñèíòåç ãë³êîãåíó åíåðãîºìíèé ïðîöåñ (çàòðà÷àºòüñÿ 2-³ ìîëåêóëè
ÀÒÔ); àâòîêàòàë³òè÷íèé (âåëèê³ ê³ëüêîñò³ â³ëüíîãî ãëþêîçî-6-ôîñôàòó
àêòè-âóþòü ãë³êîãåíñèíòåòàçó–ïðîöåñè ãë³êîãåíåçó ïîñèëþþòüñÿ; ï³äâèùåí
ê³ëüêîñò³ ÓÄÔ ³íàêòèâóþòü ãë³êîãåíñèíòåòàçó–ïðîöåñè ãë³êîãåíåçó ïðèïèíÿþòüñÿ).
Óòâîðåííÿ ãë³êîãåíó ïðîõîäèòü ç ìàêñèìàëüíîþ øâèäê³ñòüþ ÷åðåç 30–40 õâ. ï³ñëÿ
ïðèéîìó ¿æ³. Ïðîöåñè ãë³êîãåíåçó ïðîõîäÿòü ³íòåíñèâí³øå ï³ñëÿ àêòèâíî¿ ì‘ÿçåâî
ðîáîòè.
3.2 . Ãë³êîãåíîë³ç
Ïðè ì‘ÿçåâ³é ðîáîò³, ñèëüíîìó ïåðåîõîëîäæåíí³, ãîëîäóâàíí³, åìîö³éíîìó
çáóäæåíí³ â³äáóâàºòüñÿ ïîñèëåíå ñïîæèâàííÿ êë³òèíàìè ãëþêîçè êðîâ³. гâåíü
îñòàííüî¿ ïîâèíåí áóâ áè çíèæóâàòèñÿ. Ïðîòå, ó çäîðîâèõ ëþäåé ïðè ð³çíèõ
ôóíêö³îíàëüíèõ ñòàíàõ, ð³âåíü ãëþêîçè ïîñò³éíèé (120 ìã/100ñì3) ç
íåçíà÷íèìè â³äõèëåííÿìè. Ñòàá³ëüíèé ð³âåíü ãëþêîçè ó êðîâ³ çà ð³çíèõ
ôóíêö³îíàëüíèõ ñòàí³â çàáåçïå÷óºòüñÿ ïðîò³êàííÿì ïðîöåñ³â ãë³êîãåíîë³çó (ðîçïàä
ãë³êîãåíó äî ãëþêîçè). Ãë³êîãåíîë³ç ìîæå ïðîõîäèòè äâîìà øëÿõàìè: ã³äðîë³ç–ïîâ³ëüíèé,
ìàëîåôåêòèâíèé ïðîöåñ; ôîñôîðîë³ç–îñíîâíèé øëÿõ ðîçïàäó ãë³êîãåíó.
Ôîñôîðîë³ç ïðîõîäèòü çà ó÷àñòþ ôîñôîðèëàçíèõ ôåðìåíòíèõ ñèñòåì, ÿê³ íà
ïåðøîìó åòàï³ â³äùåïëþþòü â³ä ìîëåêóëè ãë³êîãåíó ê³íöåâ³ çàëèøêè ìîëåêóë ãëþêîçè
ó âèãëÿä³ ãëþêîçî-1-ôîñôàòó:
1.
CH2OH
![](/image/16236_5_36.png) ![](/image/16236_5_37.png) ![](/image/16236_5_38.png) ![](/image/16236_5_39.png) ![](/image/16236_5_40.png) ![](/image/16236_6_15.png) ![](/image/16236_7_15.png) ![](/image/16236_8_15.png) ![](/image/16236_8_16.png) ![](/image/16236_6_16.png) H
H H
(C6H10O5)n H3PO4;
(C6H10O5)n-1 OH
![](/image/16236_14_3.png) ôîñôîðèëàçà OH OH H O–Ð
H OH OH
Ãë³êîãåí Ãëþêîçî–1–ôîñôàò
2. Îñòàííÿ øâèäêî ³çîìåðèçóºòüñÿ â ãëþêîçî-6-ôîñôàò:
![](/image/16236_28_1.png) ÍÎ ÎÍ
![](/image/16236_4_9.png) CH2OH
CH2O–Ð=Î
![](/image/16236_5_41.png) ![](/image/16236_5_42.png) ![](/image/16236_5_43.png) ![](/image/16236_5_44.png) ![](/image/16236_5_45.png) ![](/image/16236_6_17.png) ![](/image/16236_7_17.png) ![](/image/16236_8_17.png) ![](/image/16236_8_18.png) ![](/image/16236_6_18.png) ![](/image/16236_7_18.png) ![](/image/16236_5_46.png) ![](/image/16236_5_47.png) ![](/image/16236_5_48.png) ![](/image/16236_5_49.png) ![](/image/16236_5_50.png) ![](/image/16236_6_19.png) ![](/image/16236_7_19.png) ![](/image/16236_8_19.png) ![](/image/16236_8_20.png) ![](/image/16236_6_20.png) H H
H H H H
![](/image/16236_19_3.png) OH
ôîñôîãëþêî³çîìåðàçà
![](/image/16236_21_3.png) ![](/image/16236_22_3.png) OH OH H O–P=O OH OH H OÍ
H OH OH
H OH
Ãëþêîçî–1–ôîñôàò Ãëþêîçî-6-ôîñôàò
3. Ãëþêîçî-6-ôîñôàò
ðîçùåïëþºòüñÿ ôîñôàòàçàìè äî â³ëüíî¿ ãëþêîçè ³ ôîñôîðíî¿ êèñëîòè:
![](/image/16236_28_2.png) ÍÎ ÎÍ
![](/image/16236_4_11.png) CH2O–Ð=Î
CH2OÍ
![](/image/16236_5_51.png) ![](/image/16236_5_52.png) ![](/image/16236_5_53.png) ![](/image/16236_5_54.png) ![](/image/16236_5_55.png) ![](/image/16236_6_21.png) ![](/image/16236_7_21.png) ![](/image/16236_8_21.png) ![](/image/16236_8_22.png) ![](/image/16236_6_22.png) ![](/image/16236_7_22.png) ![](/image/16236_5_56.png) ![](/image/16236_5_57.png) ![](/image/16236_5_58.png) ![](/image/16236_5_59.png) ![](/image/16236_5_60.png) ![](/image/16236_6_23.png) ![](/image/16236_7_23.png) ![](/image/16236_8_23.png) ![](/image/16236_8_24.png) ![](/image/16236_6_24.png) H H
H H H H
![](/image/16236_19_4.png) OH
ôîñôàòàçà + Í3ÐÎ4
![](/image/16236_21_4.png) ![](/image/16236_22_4.png) OH OH H O–P=O OH OH H OÍ
H OH OH Í2Î
H OH
Ãëþêîçî–6–ôîñôàò Ãëþêîçà
Âèñíîâîê:
ïîñò³éíà êîíöåíòðàö³ÿ ãëþêîçè ó êðîâ³ ï³äòðèìóºòüñÿ ïðîöåñàìè
ãë³êîãåíîë³çó ÿê³ çíàõîäÿòüñÿ ï³ä êîíòðîëåì åíäîêðèííî¿ ñèñòåìè: ãîðìîí
ãëþêàãîí ñòèìóëþº ïðîöåñè ðîçïàäó ãë³êîãåíó äî ãëþêîçè ó ïå÷³íö³ à àäðåíàë³í– ó
ì‘ÿçàõ òà ³íøèõ âíóòð³øí³õ îðãàíàõ. Ïðîöåñè ãë³êîãåíîë³çó ïðèñêîðþþòüñÿ ïðè
ïîñèëåí³é ì‘ÿçåâ³é ðîáîò³, ïåðåîõîëîäæåíí³, åìîö³éíîìó çáóäæåíí³. Íåçàëåæíî â³ä
åíåðãîòðàò îðãàí³çìó çàâæäè â ïå÷³íö³ çàëèøàºòüñÿ ïåâíà ê³ëüê³ñòü ãë³êîãåíó (ðåçåðâ
äëÿ ðîáîòè ãîëîâíîãî ìîçêó, ñåðöåâîãî ì‘ÿçó ³ «çàòðàâî÷íèé ãë³êîãåí»).
ϳäòðèìàííÿ ñòàíó ãîìåîñòàçó òà âèêîíàííÿ ïåâíèõ ô³çè÷íèõ âïðàâ ïðèâîäèòü
äî ïîñèëåííÿ ïðîöåñ³â ðîçïàäó ãë³êîãåíó òà îêèñëåííÿ ãëþêîçè. Îñòàííº ìîæå
â³äáóâàòèñÿ çà áåçêèñíåâèõ (àíàåðîáíèõ) óìîâ òà ó ïðèñóòíîñò³ êèñíþ (àåðîáíå
îêèñëåííÿ).
Ïî÷àòêîâîþ ñòà䳺þ îêèñëåííÿ ãëþêîçè º àíàåðîáí³ ïåðåòâîðåííÿ.
3.3 Àíàåðîáíèé ðîçïàä ãëþêîçè (ãë³êîë³ç)
Óìîâíî ïîä³ëÿþòü íà äâ³ ñòà䳿 – ï³äãîòîâ÷ó ³ îñíîâíó.
ϳäãîòîâ÷à ñòàä³ÿ ãë³êîë³çó ðîçïî÷èíàºòüñÿ ç àêòèâàö³¿ ìîëåêóëè
ãëþêîçè, ÿêà ÷åðåç ðÿä ôåðìåíòàòèâíèõ ðåàêö³é, ïåðåòâîðþºòüñÿ íà äâ³ ìîëåêóëè
ôîñôîãë³öå-ðèíîâîãî àëüäåã³äó:
![](/image/16236_2_3.png) HO OH
![](/image/16236_4_13.png) CH2OH
CH2O–P=O
![](/image/16236_5_61.png) ![](/image/16236_5_62.png) ![](/image/16236_5_63.png) ![](/image/16236_5_64.png) ![](/image/16236_5_65.png) ![](/image/16236_6_25.png) ![](/image/16236_7_25.png) ![](/image/16236_8_25.png) ![](/image/16236_8_26.png) ![](/image/16236_6_26.png) ![](/image/16236_7_26.png) ![](/image/16236_5_66.png) ![](/image/16236_5_67.png) ![](/image/16236_5_68.png) ![](/image/16236_5_69.png) ![](/image/16236_5_70.png) ![](/image/16236_6_27.png) ![](/image/16236_7_27.png) ![](/image/16236_8_27.png) ![](/image/16236_8_28.png) ![](/image/16236_6_28.png) H H
H ÀÒÔ ÀÄÔ H H
H
![](/image/16236_30_1.png)
OH OH H OH ãåêñîê³íàçà OH
OH H OH
H OH H OH
Ãëþêîçà Ãëþêîçî–6–ôîñôàò
![](/image/16236_12_2.png) HO OH
![](/image/16236_31_1.png) ![](/image/16236_32_1.png) ![](/image/16236_33_1.png) CH2O–P=O ÍÎ ÎÍ
![](/image/16236_34_1.png) ![](/image/16236_35_1.png) ![](/image/16236_36_1.png) ![](/image/16236_5_71.png) ![](/image/16236_5_72.png) ![](/image/16236_5_73.png) ![](/image/16236_5_74.png) ![](/image/16236_5_75.png) ![](/image/16236_6_29.png) ![](/image/16236_7_29.png) ![](/image/16236_8_29.png) ![](/image/16236_8_30.png) ![](/image/16236_6_30.png) H H
H O=P–O–CH2 OH
![](/image/16236_37_1.png) ![](/image/16236_38_1.png) ôîñôîãëþêî³çîìåðàçà H HO
![](/image/16236_40_1.png) ![](/image/16236_41_1.png) ![](/image/16236_42_1.png) OH OH
H OH
H OH H
OH H CH2OH
Ãëþêîçî–6–ôîñôàò Ôðóêòîçî–6–ôîñôàò
![](/image/16236_43_1.png) O
![](/image/16236_44_1.png) ![](/image/16236_45_1.png) ![](/image/16236_46_1.png) ![](/image/16236_47_1.png) HO–P–OH HO OH
![](/image/16236_48_1.png) ![](/image/16236_49_1.png) ![](/image/16236_50_1.png) ![](/image/16236_34_2.png) ![](/image/16236_35_2.png) ![](/image/16236_36_2.png) ![](/image/16236_5_76.png) ![](/image/16236_5_77.png) ![](/image/16236_5_78.png) ![](/image/16236_6_31.png) ![](/image/16236_7_31.png) O–CH2
OH O=P–O–CH2 OH
![](/image/16236_51_1.png) ![](/image/16236_38_2.png) ôîñôîôðóêòîê³íàçà H HO
![](/image/16236_52_1.png) ![](/image/16236_53_1.png) ![](/image/16236_54_1.png) ![](/image/16236_40_2.png) ![](/image/16236_41_2.png) H H
OH HO OH
OH H CH2ÎH H OH H CH2O–P=O
Ôðóêòîçî–6–ôîñôàò Ôðóêòîçî–1,6–äèôîñôàò
![](/image/16236_43_3.png) O
![](/image/16236_55_1.png) ![](/image/16236_46_2.png) HO–P–OH OH
![](/image/16236_56_1.png) ![](/image/16236_56_2.png) ![](/image/16236_57_1.png) ![](/image/16236_58_1.png) ![](/image/16236_56_3.png) ![](/image/16236_56_4.png) ![](/image/16236_56_5.png) ![](/image/16236_56_6.png) ![](/image/16236_56_7.png) ![](/image/16236_56_8.png) ![](/image/16236_48_2.png) ![](/image/16236_49_2.png) ![](/image/16236_50_2.png) ![](/image/16236_5_79.png) ![](/image/16236_5_80.png) ![](/image/16236_5_81.png) ![](/image/16236_6_32.png) ![](/image/16236_7_32.png) O–CH2
OH CH2–OP=O OH
àëüäîëàçà C=O OH + CH2–OP=O
![](/image/16236_52_2.png) ![](/image/16236_53_2.png) H H OH HO OH CH2–OH CH=O
OH
OH H CH2ΖP=O CH2–OH
Ôðóêòîçî–1,6–äèôîñôàò Äèîêñ³àöåòîíôîñôàò
![](/image/16236_59_1.png) OH
OH
![](/image/16236_61_1.png) CH2–OP=O CH2–O–P=O
CH=O OH ³çîìåðàçà CH–OH OH
CH2–OH C=O
H
Äèîêñ³àöåòîíôîñôàò Ôîñôîãë³öåðèíîâèé
àëüäåã³ä
Îòæå:
ï³äãîòîâ÷à ñòàä³ÿ ãë³êîë³çó ðåãóëþºòüñÿ äâîìà àëîñòåðè÷íèìè ôàêòîðàìè: ãåêñîê³íàçíèìè òà ôîñôîôðóêòîê³íàçíèìè ôåðìåíòíèìè ñèñòåìàìè, ÿê³ ïðè íèçüêèõ êîíöåíòðàö³ÿõ ÀÒÔ ó êë³òèí³ çàïóñêàþòü ïðîöåñè
ãë³êîë³çó.  õîä³ ï³äãî-òîâ÷î¿ ñòàä
ãë³êîë³çó âèòðà÷àºòüñÿ äâ³ ìîëåêóëè ÀÒÔ.
Îñíîâíà ñòàä³ÿ ãë³êîë³çó – ñóêóïí³ñòü ôåðìåíòàòèâíèõ ðåàêö³é ïî ïåðåòâîðåííþ
ôîñôîãë³öåðèíîâîãî àëüäåã³äó äî ï³ðîâèíîãðàäíî¿ ÷è ìîëî÷íî¿ êèñëîòè.
1. Îêèñëåííÿ ÔÃÀ. äî ôîñôîãë³öåðèíîâî¿ êèñëîòè çà ó÷àñòþ ôåðìåíòó
ãë³öåð-àëüäåã³ä–3–ôîñôàòäåã³äðîãåíàçè (àêòèâíà ÷àñòèíà êîôåðìåíòó – ò³îëîâà
ãðóïà òà îêèñëåíà ôîðìà ÍÀÄ+):
![](/image/16236_65_1.png) ![](/image/16236_66_1.png) ![](/image/16236_67_1.png) ![](/image/16236_68_1.png) ![](/image/16236_69_1.png) ![](/image/16236_56_10.png) CH=O H–C–OH Ñ=Î
CH–OH SF
SF
![](/image/16236_70_1.png) ![](/image/16236_71_1.png) ![](/image/16236_72_1.png) ![](/image/16236_73_1.png) CH2–O–P=O + HS–F CH–OH ÍÀÄ+ ÍÀÄÍÍ CH–OH
![](/image/16236_75_1.png) ![](/image/16236_76_1.png) ![](/image/16236_77_1.png) ![](/image/16236_78_1.png) HO OH CH2–O–P=O CH2–O–P=O
HO OH
HO OH![](/image/16236_56_12.png)
ìàêðîåðã³÷íèé
çâ‘ÿçîê
![](/image/16236_80_1.png) ![](/image/16236_81_1.png) ![](/image/16236_82_1.png) ![](/image/16236_83_1.png) ![](/image/16236_84_1.png) C=O C=O OH ÑOOH
![](/image/16236_86_1.png) ![](/image/16236_87_1.png) SF
O–P=O CH–OH
![](/image/16236_89_1.png) ![](/image/16236_90_1.png) ![](/image/16236_91_1.png) ![](/image/16236_92_1.png) ![](/image/16236_93_1.png) CH–OH +
H3PO4 OH ÀÄÔ ÀÒÔ CH2–O–P=O
![](/image/16236_94_1.png) ![](/image/16236_26_2.png) ![](/image/16236_95_1.png) ![](/image/16236_96_1.png) CH2–O–P=O
HS–F CH–OH OH HO OH
HO OH CH2–O–P=O
OH
1,3-äèôîñôîãë³öåðèíîâà
êèñëîòà (ÔÃÊ) 3-ÔÃÊ
![](/image/16236_99_1.png) ![](/image/16236_100_1.png) ![](/image/16236_100_2.png) ÑOOH COOH
OH
![](/image/16236_101_1.png) ![](/image/16236_102_1.png) CH–OH ôîñôîãë³öåðîìóòàçà CH–O–P=O
![](/image/16236_103_1.png) CH2–O–P=O CH2–OH
OH
HO OH
3–ôîñôîãë³öåðèíîâà êèñëîòà 2–ôîñôîãë³öåðèíîâà
êèñëîòà
2. Ïåðåòâîðåííÿ ôîñôîãë³öåðèíîâî¿ êèñëîòè äî ï³ðîâèíîãðàäíî
÷è ìîëî÷íî¿ êèñëîòè:
![](/image/16236_100_4.png) ![](/image/16236_104_1.png) ![](/image/16236_100_5.png) ![](/image/16236_100_6.png) ![](/image/16236_100_7.png) ![](/image/16236_105_1.png) COOH OH COOH OH COOH
![](/image/16236_106_1.png) ![](/image/16236_106_2.png) ![](/image/16236_106_3.png) ![](/image/16236_106_4.png) ![](/image/16236_107_1.png) ![](/image/16236_108_1.png) ![](/image/16236_109_1.png) ![](/image/16236_110_1.png) ![](/image/16236_111_1.png) CH–O–P=O åíîëàçà
(–Í2Î) C–O–P=O ÀÄÔ ÀÒÔ C=O
CH2–OH OH CH2 OH CH3
2–ÔÃÊ 2–ôîñôîåíîëï³ðîâèíîãðàäíà êèñëîòà (ÏÂÊ) ÏÂÊ
![](/image/16236_113_1.png) COOH ÑÎÎÍ
![](/image/16236_115_1.png) ![](/image/16236_116_1.png) C=O ÍÀÄ
ÍÍ ÍÀÄ+ ÑÍ–ÎÍ
CH3 ÑÍ3 ÏÂÊ ìîëî÷íà
êèñëîòà
Îòæå:
â õîä³ ãë³êîë³çó âèä³ëÿºòüñÿ 6–7 % ïîòåíö³àëüíî¿ åíåð㳿 ãëþêîçè ó
âèãëÿä³ 2 ìîëåêóë ÀÒÔ. ʳíöåâèìè ïðîäóêòàìè ãë³êîë³òè÷íèõ ïðîöåñ³â îêèñëåííÿ
âóãëå-âîä³â º ï³ðîâèíîãðàäíà êèñëîòà (íîðìàëüí
ô³ç³îëîã³÷í³ óìîâè) ÷è ìîëî÷íà êèñ-ëîòà (çà íåäîñòàòíüî
ê³ëüêîñò³ êèñíþ). Îñòàííÿ, ÿê ñèëüíà îðãàí³÷íà êèñëîòà, «çàêèñëþþ÷è» ñåðåäîâèùå
íã³áóº àêòèâí³ñòü àëîñòåðè÷íèõ ôàêòîð³â, çàïîá³ãà-þ÷è ïåðåâòîì³.
Ó ìîëî÷í³é ÷è ï³ðîâèíîãðàäí³é êèñëîò³ ùå ì³ñòèòüñÿ äî 93 % ïîòåíö³éíî
åíåð㳿 ãëþêîçè, ÿêà âèä³ëÿºòüñÿ â çàêëþ÷í³é, àåðîáí³é ôàç³ ïåðåòâîðåííÿ
âóãëåâîä³â. Àåðîáíà ôàçà õàðàêòåðèçóºòüñÿ ðîçïàäîì ïðîäóêò³â ãë³êîë³çó
(ìîëî÷íî¿ êèñëîòè ÷è ÏÂÊ) äî âóãëåêèñëîãî ãàçó ³ âîäè ç âèä³ëåííÿì âåëèêî
ê³ëüêîñò³ åíåð㳿. ʳí-öåâèì àêöåïòîðîì åëåêòðîí³â
ïðîòîí³â º òêàíèííèé êèñåíü. Ñêëàäàºòüñÿ ç ï³ä-ãîòîâ÷î
ñòà䳿 ³ öèêëó òðèêàðáîíîâèõ êèñëîò.
3.4 Àåðîáíîãî ïåðåòâîðåííÿ âóãëåâîä³â
ϳäãîòîâ÷à ñòàä³ÿ àåðîáíîãî ïåðåòâîðåííÿ âóãëåâîä³â
õàðàêòåðèçóºòüñÿ ïåðåòâî-ðåííÿì ÏÂÊ ÷è ìîëî÷íî¿ êèñëîòè â
àêòèâíó ôîðìó àöåòèë–ÊîÀ. Íà äàíîìó åòàï³ âèä³ëÿºòüñÿ áëèçüêî 7 % åíåð㳿 ùî
àêóìóëüîâàíà ó â³äïîâ³äíèõ ìåòàáîë³òàõ:
CO2
![](/image/16236_119_1.png) ![](/image/16236_107_2.png) COOH ÑÎÎÍ ÍÀÄ+ ÍÀÄÍ2
![](/image/16236_120_1.png) ![](/image/16236_70_4.png) ![](/image/16236_60_2.png) CH–OH ÍÀÄ+ ÍÀÄÍÍ Ñ=Î HS–ÊîÀ ÑÍ3–Ñ=Î
![](/image/16236_122_1.png) CH3 ëàêòàòäåã³äðîãåíàçà ÑÍ3
ï³ðóâàòäåêàðáîêñèëàçà S–KoA
Öèêë òðèêàðáîíîâèõ êèñëîò – ñóêóïí³ñòü
ôåðìåíòàòèâíèõ ðåàêö³é çà ó÷àñòþ òðèêàðáîíîâàíèõ êèñëîò. Âîíè ðîçïî÷èíàþòüñÿ
âçàºìî䳺þ àöåòèë–ÊîÀ ç ùàâëåâî-îöòîâîþ êèñëîòîþ ³ çàê³í÷óþòüñÿ ðîçïàäîì
ÿáëó÷íî¿ êèñëîòè äî ùàâëåâî-îöòîâî¿ êèñëîòè.  õîä³ äàíèõ ðåàêö³é â³äíîâëþºòüñÿ
îêèñëåí³ ôîðìè äåã³äðîãåíàçíèõ ôåðìåíòíèõ ñèñòåì òà âèä³ëÿºòüñÿ âóãëåêèñëèé
ãàç:
![](/image/16236_124_1.png) ![](/image/16236_125_1.png) ![](/image/16236_126_1.png) ![](/image/16236_87_2.png) ![](/image/16236_88_2.png) C=O O=C–COOH C=O
![](/image/16236_128_1.png) SKoA + CH2 öèòðàòñèíòåòàçà
SKoA H–OH – HS–KoA
![](/image/16236_129_1.png) CH3 COOH CH2 àêîí³òàòã³äðàçà
HO
C–COOH
CH2
COOH
àöåòèë–ÊîÀ ùàâëåâî-îöòîâà êèñëîòà öèòðèë–ÊîÀ
![](/image/16236_44_2.png) ![](/image/16236_44_3.png) ![](/image/16236_44_4.png) ![](/image/16236_44_5.png) ![](/image/16236_44_6.png) ![](/image/16236_44_7.png) ![](/image/16236_44_8.png) ![](/image/16236_44_9.png) ![](/image/16236_44_10.png) ![](/image/16236_44_11.png) ![](/image/16236_44_12.png) ![](/image/16236_44_13.png) COOH COOH COOH
CH2 CH2 CH2
![](/image/16236_132_1.png) ![](/image/16236_133_1.png) ![](/image/16236_134_1.png) HO–C–COOH – H2O C–COOH +H–OH CH–COOH ÍÀÄ ÍÀÄÍÍ
CH2 CH
HO–CH ³çîöèòðàò-–
COOH COOH COOH äåã³äðîãåíàçà
ëèìîííà êèñëîòà öèñ-àêîí³òîâà ³çîëèìîííà
![](/image/16236_136_1.png)
CO2 – CO2
COOH COOH COOH
![](/image/16236_138_1.png) ![](/image/16236_44_15.png) ![](/image/16236_44_16.png) ![](/image/16236_44_17.png) ![](/image/16236_44_18.png) CH2 CH2 +ÍÀÄ+ ÍÀÄÍÍ CH2
![](/image/16236_140_1.png) ![](/image/16236_141_1.png) CH–COOH CH2 +ÍS–F –CO2 CH2
![](/image/16236_143_1.png) ![](/image/16236_144_1.png) C=O C=O –êåòîãëóòàðàò– C=O
![](/image/16236_44_19.png) ![](/image/16236_44_20.png) ![](/image/16236_44_21.png) ![](/image/16236_44_22.png) ![](/image/16236_44_23.png) ![](/image/16236_44_24.png) COOH COOH äåã³äðîãåíàçà S–KoA
ùàâëåâî–ÿíòàðíà –êåòîãëóòàðîâà ñóêöèí³ë–ÊîÀ
![](/image/16236_147_1.png) ![](/image/16236_147_2.png) ![](/image/16236_147_3.png) ![](/image/16236_147_4.png) ![](/image/16236_147_5.png) ![](/image/16236_147_6.png) ![](/image/16236_147_7.png) ![](/image/16236_147_8.png) ![](/image/16236_147_9.png) ![](/image/16236_147_10.png) ![](/image/16236_147_11.png) ![](/image/16236_147_12.png) COOH ÑOOH COOH
CH2 CH
CH2
![](/image/16236_148_1.png) ![](/image/16236_149_1.png) ![](/image/16236_150_1.png) ÃÄÔ ÃÒÔ CH2 ÔÀÄ+ ÔÀÄÍÍ CH H–OH CH–OH ÍÀÄ+ ÍÀÄÍÍ
![](/image/16236_152_1.png) COOH COOH COOH
ÿíòàðíà êèñëîòà ôóìàðîâà ÿáëó÷íà
êèñëîòà
ÑOOH
CH2
C=O
COOH
ùàâëåâî-îöòîâà êèñëîòà
Îòæå:
åíåðãåòè÷íèé åôåêò ï³äãîòîâ÷î¿õ ñòà䳿 àåðîáíîãî îêèñëåííÿ ñòàíîâèòü 6 ìîëåêóë ÀÒÔ (äâ³ â³äíîâëåí³ ôîðìè ÍÀÄ-çàëåæíèõ ôåðìåíò³â); åíåðãåòè÷íèé åôåêò
öèêëó Êðåáñà {3 ÍÀÄÍÍ, 1ÔÀÄÍÍ, 1ÃÒÔ}=12 ìîëåêóë ÀÒÔ.
Ïîñê³ëüêè îäíà ìîëåêóëà ãëþêîçè ðîçêëàäàºòüñÿ íà äâ³ ìîëåêóëè ôîñôîãë³öåðè-íîâîãî àëüäåã³äó, òî çàãàëüíèé åíåðãåòè÷íèé
åôåêò àåðîáíîãî îêèñëåííÿ ãëþêîçè ñòàíîâèòü 18 2=36 ìîëåêóë ÀÒÔ, à âðàõîâóþ÷è äâ³ ìîëåêóëè ÀÒÔ ãë³êîë³òè÷-íîãî
îêèñëåííÿ –38 ìîëåêóë ÀÒÔ.
 38-è ìîëåêóëàõ ÀÒÔ àêóìóëþºòüñÿ ò³ëüêè 50 % ïîòåíö³éíî¿ åíåð㳿 ãëþêîçè ( êÄæ)
à ³íø³ 50 % âèä³ëÿþòüñÿ ó âèãëÿä³ òåïëà.
Ïèòàííÿ äëÿ ñàìîêîíòðîëþ
1. Ðîçêðèòè ñóòü ôåðìåíòàòèâíîãî ïåðåòâîðåííÿ âóãëåâîä³â ó õàð÷îâîìó
òðàêò³ ëþäèíè.
2. Ïîøèðåííÿ âóãëåâîä³â ó îðãàí³çì³ ëþäèíè. Ô³ç³îëîã³÷íå çíà÷åííÿ
ãëþêîçè.
3. Ñèíòåç ãë³êîãåíó ç ãëþêîçè (ãë³êîãåíåç). Õ³ì³çì ïðîöåñó, ôàêòîðè
ðåãóëÿö³¿.
4. Ðîçïàä ãë³êîãåíó äî ãëþêîçè (ãë³êîãåíîë³ç). Õ³ì³çì ïðîöåñó, ôàêòîðè
ðåãóëÿö³¿.
5. Õàðàêòåðèñòèêà ï³äãîòîâ÷î¿ òà îñíîâíî¿ ñòàä³é àíàåðîáíîãî ðîçïàäó
(ãë³êîë³ç). Õ³ì³çì ïðîöåñó, ôàêòîðè ðåãóëÿö³¿, åíåðãåòè÷í³ åôåêòè.
6. Õàðàêòåðèñòèêà ï³äãîòîâ÷î¿ òà îñíîâíî¿ ñòàä³é àåðîáíîãî ðîçïàäó
âóãëåâîä³â. Õ³ì³çì ïðîöåñó, ôàêòîðè ðåãóëÿö³¿, åíåðãåòè÷í³ åôåêòè.
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