1 lehninger principles of biochemistry fourth edition chapter 20: carbohydrate biosynthesis in...
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Lehninger Principles of BiochemistryFourth Edition
Chapter 20:Carbohydrate Biosynthesis in
Plantsand Bacteria
Copyright © 2004 by W. H. Freeman & Company
David L. Nelson and Michael M. Cox
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Ensímið rubisco
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Nokkur orð og hugtök (án ábyrgðar!)
Plastid: Plastíð (ekki sama og plasmíð) geyma forða eða litarefni í plöntum, skiptast í:
Chloroplasts: grænukorn, innihalda blaðgrænu (chlorophyll)Chromoplasts: litkorn, innihalda rauð eða gul litarefniAmyloplasts: innihalda forðanæringu en engin litarefni,
kallast líka leucoplasts (hvítkorn)
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Joðlitun fyrir sterkju í sterkjukornum
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Hvað eru ljósóháð hvörf í plöntum og hvar gerast þau?
Í ljósóháðum hvörfum (light-independent reactions, dark reactions) tengist CO2 við sykrur
Þetta gerist í grænukornsmerg
NADPH og ATP sem ljóskerfin mynda eru notuð til efnasmíðarinnar
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Hvað gerist í Calvin-hringrás, hvernig og hverjar eru afurðirnir?
CO2 kemur inn í hringinn og hvarfast við virkjað pentósafosfat (ríbúlósa-1,5-bisfosfat)
Ríbúlósa-1,5-bisfosfat endurnýjast í lok hringsins, sbr. oxalóasetat í sítrónusýruhring
Ensímið sem hvetur hvörfin er ríbúlósa-1,5-bisfosfatkarboxýlasi/oxýgenasi (“rubisco”)
Rubisco er algengasta ensím á jörðu og líklega algengasta próteinið (16% grænukornspróteina)
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Calvin-hringur - yfirlit
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Calvin-hringur
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Ensímið rubisco
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Rubisco
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Mg2+ í rubisco
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Fyrsta myndefnið er óstöðug, greinótt -ketósýraFrá henni myndast tvær sameindir tríósafosfats (3-fosfóglýserat)
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Hvarfagangur rubisco
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Stýrilsameind rubisco
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Virkjun rubisco
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3-fosfóglýserat hvarfast síðan í glýseraldehýð-3-fosfat í tveimur skrefum
Í fyrra skrefinu er þörf fyrir ATP en NADPH í hinu seinna
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Afoxunarhvörf Calvin-hrings
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Afdrif tríósafosfata:Hluti þeirra er notaður til að mynda sykrur (sterkju, súkrósa), sem eru teknar út úr hringnum Hluti þeirra er notaður til að endurmynda ríbúlósa-1,5-bisfosfat í ferli sem líkist pentósaferli
Uppstokkunarskref pentósaferlis eru gagnhverf, en Calvin-hringur er ógagnhverfur
Í Calvin-hring er þörf fyrir ATP til að fosfórýlera sykrurNokkur skref eru ógagnhverf, þau sem eru hvött af rubisco og fosfatösum svo og myndun ríbúlósa-1,5-bisfosfats
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Uppstokkunarskref Calvin-hrings - yfirlit
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Transketólasahvött hvörf í Calvin-hring
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Þíamínpýrófosfat er kófaktor í Calvin-hring
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Endurmyndun ríbúlósa-1,5-bisfosfats er ógagnhverf
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Einföld jafna ljóstillífunar er:
6 CO2 + 6 H2O C6H12O6 + 6O2
Heildarjafna Calvin-hrings er (fremur til að átta sig á heildarmyndinni en að læra hana!):
6 CO2 + 12 H2O + 18 ATP + 12 NADPH + 12 H+ C6H12O6 + 18 ADP + 18 Pi + 12 NADP+
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Calvin-hringur - yfirlit
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Flutningur afurða út í frymi
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Flutningur afoxunarvalds (NADPH) og ATP í frymi
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Uppruni ATP og NADPH
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FBPasi virkjast við basískar aðstæður
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Ljósháð afoxunarhvörf virkja sum ensím
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Hvað er ljósöndun (photorespiration)?
Rubisco er ekki fullkomið ensímÞað getur einnig hvarfast við súrefni, er oxýgenasi eins og nafnið gefur til kynna
Með hækkandi hitastigi minnkar sækni rubisco í CO2
Hlutfall O2/CO2 í lausn eykst með hækkandi hitastigi
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Ljósöndun - oxýgenasavirkni rubsico
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Við oxýgenasahvörfin myndast 3-fosfóglýserat
Einnig myndast 2-fosfóglýkólat sem er unnt að endurvinna en það er orkulega dýrtLjósöndun er því orkulega og efnislega óhagstæðÞetta er vandamál fyrir hitabeltisjurtir Reynt hefur verið að “endurbæta” rubisco með erfðatækni, en ekki tekist
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Ljósöndun er orkulega og efnislega óhagstæð
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Glýkólatbraut endurnýtir afurðir ljósöndunar og er dýr
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Glýsíndekarboxýlasi hefur flókinn hvarfagang
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Hvernig sneiða hitabeltisjurtir (maís, sykurreyr, kaktusar, ananas) hjá ljósöndun?
Þær innlima CO2 í 3ja kolefnisatóma sýrur og mynda malat (4 C-atóm)Í venjulegum plöntum í tempraða beltinu myndast 3ja kolefnisatóma sýrur (C3-plöntur)Í öllum plöntum eru mesophyll cells (blaðholdsfrumur) nálægt yfirborði plöntunnar
Blaðhold er sá plöntuvefur sem inniheldur grænukorn
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Í maís, sykurreyr og fleiri plöntum (C4-plöntur) er blaðholdið óvenjulegt þar sem það inniheldur ekki rubisco
Malatið er smíðað í blaðholdi, en geymt í sérstökum frumuhlutum – aðskilið í rúmi
Fyrir neðan blaðholdið eru strengslíðurfrumur (bundle sheath cells) í öllum plöntum
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Í C4-plöntum innihalda strengslíðurfrumur rubisco, aðrar plöntur ekkiÍ C4-plöntum er malat flutt úr blaðholdsfrumum og inn í strengslíðurfrumur
Í strengslíðurfrumum tapar malatið CO2, sem rubisco innlimar í ríbúlósa-1,5-bisfosfat
C3-sýran sem myndast er síðan flutt úr strengslíðurfrumum í blaðholdsfrumur
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Maís og sykurreyr geyma malat í sérsökum frumuhlutum
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Succulent plants: Þykkblöðungar, safajurtir
Kaktusar og ananas loka loftaugum (stomata) á daginn til að varðveita vatn
Á nóttunni opnast loftaugun, CO2 kemst inn og er innlimað í 3ja kolefnisatóma sýrur og malat myndast (4 kolefnisatóm) – ferlið er aðskilið í tíma
Að morgni þegar ljóstillífun hefst, losnar CO2 frá malati, og það dregur úr ljósöndun
Þessar plöntur kallast CAM-plöntur (Crassulacean Acid Metabolism)Crassulacea eru plöntur af hnoðraætt
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Bæði þessi ferli kosta meira ATPÍ staðinn fyrir að það kosti 3 ATP að innlima eina CO2-sameind, kostar það 5 ATP
Í venjulegri C3-plöntu eykst ljósöndunin með hækkandi hitastigi og plantan vex hægar
Með hækkandi hitastigi standa C4-plönturnar betur að vígi þrátt fyrir aukna ATP-þörf
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Smíð sterkju, súkrósa og sellúlósa
Smíð sterkju er hvött af sterkjusýnþasa, ADP-glúkósi er virkjuð glúkósaeining (ekki UDPGlc)
Vaxandi keðjum er bætt til skiptis á tvö set á ensíminu, að lokum losnar sterkjan frá
Sterkjusmíð er stýrt með myndun upphafsefnis sterkjusmíðar (ADPGlc)
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Smíð sterkju
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Sterkjusmíð er stýrt með myndun upphafsefnis sterkjusmíðar (ADPGlc)
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Súkrósi er smíðaður frá UDP-glúkósa og frúktósa-6-fosfati í tveimur skrefumFyrsta afurðin er súkrósa-6-fosfat, fosfatið losnar frá í öðru skrefi og súkrósi myndsast
Frúktósa-2,6-bisfosfat er stýrilsameind súkrósasmíðarSúkrósasmíð er stýrt með ábót og brottnámi fosfats á ensím
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Smíð súkrósa
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Frúktósa-2,6-bisfosfat er stýrilsameind súkrósasmíðar
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Súkrósasmíð er stýrt með ábót og brottnámi fosfats á ensím
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Sellúlósi er línuleg stoðfjölsykra og hómópólýsakkaríðÍ sellúlósa eru tengin milli glúkósaleifa -1,4-tengi
Sellúlósi hefur þráðlaga byggingu sem er gerð stöðug af vetnistengjum milli sameinda
Spendýr framleiða ekki meltingarensím sem rjúfa -1,4-tengi
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Bygging sellúlósa
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Sellúlósasmíð gerist í komplexumHimnubundinn plöntusteri (líklega sitosterol) er “primer” í sellúlósasmíð
Sykrueiningu (UDPGlc) er bætt á hýdroxýlhóp sterans á innra borði frumuhimnu
Þegar ákveðinni keðjulengd er náð, “flippar” einingin yfir himnunaÞar er hún framlengd og losuð frá
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Sellúlósasmíð gerist í komplexum
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Himnubundinn plöntusteri er “primer” í sellúlósasmíð
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Frumuveggir baktería
Frumuveggir baktería eru að stofni til línuleg fjölliða með endurtekinni tvísykrueiningu
Fjölsykrukeðjurnar eru þvertengdar með litlum peptíðakeðjum og mynda þéttriðið net
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Smíð frumuveggja í bakteríum
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Penisillín hindrar tengingu annarrar peptíðakeðjunnar við sykrukeðjurnar með því að mynda hvarfastigshermi (transition-state analog) við peptíðabút
Sumar örverur gera penisillín óvirkt
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Penisillín hindrar skref í smíð frumuveggjar baktería
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Sumar örverur gera penisillín óvirkt
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Ensímið lysozyme finnst m. a. í ýmsu seyti: tárum, munnvatni og broddmjólk, einnig eggjahvítu
Ensímið hvetur vatnsrof annars glýkósíðtengjanna í sykrukeðjunni, þannig að tvísykruafleiða myndast og frumuveggurinn rofnar
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Í kímplöntum koma sykrur frá fitum (glýoxýlathringur)
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Afdrif glýserólgrindar - fer í nýmyndun glúkósa
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Plöntur hafa birgðir af hexósafosfötum, tríósafosfötum og pentósafosfötum
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