Hoa Sinh

TRNG AI HOC A LAT
GIAO TRNH
HOA SINH HOC

GS.TS. MAI XUAN LNG

2001
Hoa sinh hoc - 1 -
MUC LUC
MUC LUC................................................................................................................. - 1 -
M AU................................................................................................................... - 5 -
CHNG 1. AMINOACID VA PROTEIN ............................................................ - 10 -
I. AMINOACID. .................................................................................................. - 10 -
1. Cau tao......................................................................................................... - 11 -
2. Hoat tnh quang hoc. .................................................................................... - 13 -
3. Tnh chat lng tnh. .................................................................................... - 14 -
4. Cac phan ng hoa hoc ac trng. ................................................................. - 16 -
II. PEPTIDE......................................................................................................... - 19 -
III. TNH CHAT CUA LIEN KET PEPTIDE...................................................... - 21 -
IV. CAC LIEN KET TH CAP TRONG PHAN T PROTEIN. ........................ - 21 -
V. CAU TRUC CUA PROTEIN.......................................................................... - 23 -
1.Cau truc bac mot........................................................................................... - 23 -
2. Cau truc bac hai. .......................................................................................... - 23 -
3. Cau truc bac ba. ........................................................................................... - 25 -
4. Cau truc bac bon. ......................................................................................... - 25 -
VI. TNH CHAT CUA PROTEIN. ...................................................................... - 26 -
1. Tnh chat lng tnh. .................................................................................... - 26 -
2. Hoat tnh quang hoc. .................................................................................... - 26 -
3. Tnh hydrate-hoa. ........................................................................................ - 26 -
4. S bien tnh cua protein. .............................................................................. - 27 -
5. Cac phan ng mau ac trng. ...................................................................... - 28 -
6. Hoat tnh va chc nang sinh hoc cua protein. .............................................. - 28 -
VII. PHAN LOAI PROTEIN. .............................................................................. - 29 -
VIII. PHAN GIAI PROTEIN. .............................................................................. - 31 -
IX. PHAN GIAI AMINOACID. .......................................................................... - 32 -
1.Chuyen amin hoa. ......................................................................................... - 32 -
2. Desamin hoa. ............................................................................................... - 33 -
3. Decarboxyl hoa............................................................................................ - 34 -
4. So phan cua ammoniac va chu trnh urea. ................................................... - 34 -
5. D hoa aminoacid va chu trnh acid tricarboxylic. ....................................... - 35 -
X. SINH TONG HP AMINOACID................................................................... - 36 -
1.Kh nitrate va co nh nit. .......................................................................... - 36 -
2. Amin hoa kh............................................................................................... - 37 -
3. Tong hp cac aminoacid th cap. ................................................................ - 37 -
XI. SINH TONG HP PROTEIN........................................................................ - 38 -
1. Cac yeu to can thiet cho sinh tong hp protein va cac giai oan cua qua trnh
nay. .................................................................................................................. - 38 -
2. ieu hoa sinh tong hp protein; mo hnh operon va ly thuyet ieu hoa cua
Jacob va Monod............................................................................................... - 41 -
GS.TS. Mai Xuan Lng Khoa Sinh hoc

Hoa sinh hoc - 2 -
CHNG 2. EMZYME.......................................................................................... - 45 -
I. CAC BIEU THC DUNG TRONG ENZYME HOC....................................... - 45 -
II. BAN CHAT HOA HOC CUA ENZYME. ...................................................... - 45 -
III. ONG HOC CUA CAC PHAN NG ENZYME. ......................................... - 47 -
IV. ANH HNG CUA pH LEN HOAT TNH ENZYME. ............................... - 50 -
V. ANH HNG CUA NHIET O LEN HOAT TNH ENZYME. .................. - 51 -
VI. HOAT HOA ENZYME. ................................................................................ - 51 -
VII. C CHE ENZYME. ..................................................................................... - 52 -
VIII. TNH AC HIEU CUA ENZYME. ............................................................ - 57 -
IX. DANH PHAP VA PHAN LOAI ENZYME. .................................................. - 58 -
X. HE THONG MULTIENZYM VA VAI TRO CUA ENZYME IEU HOA.... - 59 -
XI. ISOENZYME................................................................................................ - 61 -
CHNG 3. KHAI NIEM CHUNG VE TRAO OI CHAT .................................. - 63 -
I. ONG HOA VA D HOA. ............................................................................... - 63 -
II. CAC HNH THC VAN CHUYEN NANG LNG TRONG TRAO OI
CHAT. ................................................................................................................. - 65 -
III. NANG LNG SINH HOC VA CHU TRNH ATP..................................... - 66 -
IV. VAN CHUYEN NANG LNG TRONG CAC PHAN NG OXY HOA KH
............................................................................................................................. - 70 -
CHNG 4. GLUCID........................................................................................... - 73 -
I. MONOSACHARIDE (MONOSE) ................................................................... - 73 -
1. cau tao.......................................................................................................... - 73 -
2. Tnh chat hoa hoc......................................................................................... - 77 -
II. OLIGOSACCHARIDE. .................................................................................. - 80 -
1.Disacchride. .................................................................................................. - 80 -
2.Trisaccharide. ............................................................................................... - 81 -
3.Tetrasaccharide. ........................................................................................... - 81 -
III. POLYSACCHARIDE (POLYOSE)............................................................... - 81 -
1.Homopolisaccharide. .................................................................................... - 82 -
2.Heteropolysaccharide. .................................................................................. - 85 -
IV. PHAN GIAI POLYSACCHARIDE. .............................................................. - 88 -
1.Phan giai tinh bot va glycogen...................................................................... - 88 -
2.Phan giai cac polysaccharide khac. .............................................................. - 90 -
V. CHUYEN HOA TNG HO GIA CAC MONOSE. ................................... - 90 -
1.Trao oi (van chuyen) cac nhom glycosyl cua glycosylphosphate:.............. - 90 -
2.Epimer hoa: .................................................................................................. - 90 -
3.Oxy hoa hexose va decarboxyl hoa thanh pentose: ..................................... - 91 -
VI. GLYCOLYS. ................................................................................................. - 91 -
VII. CHU TRNH PENTOSOPHOSPHATE ....................................................... - 95 -
VIII. OXY HOA HIEU KH GLUCID. ............................................................... - 96 -
1.Decarboxyl hoa oxy hoa acid pyruvic. ......................................................... - 96 -

Hoa sinh hoc - 3 -
2. Chu trnh acid tricarboxylic (Chu trnh Krebs) ............................................ - 97 -
3. Y ngha cua chu trnh acid tricarboxylic. ..................................................... - 98 -
4. Cac phan ng bu ap. .................................................................................. - 99 -
IX. PHOSPHORYL HOA OXY HOA. ................................................................ - 99 -
X. QUANG HP ............................................................................................... - 103 -
1. Phng trnh tong quat cua quang hp...................................................... - 103 -
2. Khai niem ve tch chat hai giai oan cua quang hp. ................................ - 104 -
3. Vai tro cua nang lng anh sang oi vi quang hp. ................................. - 105 -
4. C s cau truc cua quang phosphoryl-hoa. ............................................... - 111 -
5. Co nh CO
2
trong pha toi cua quang hp................................................. - 113 -
CHNG 5. LIPID.............................................................................................. - 117 -
I. ACID BEO. .................................................................................................... - 117 -
II.CAC ESTER CUA GLYCEROL. .................................................................. - 119 -
1.Lipid trung tnh. .......................................................................................... - 119 -
2.Phosphatide................................................................................................. - 122 -
3.Glycerogalactolipid va glycerosulfolipid.................................................... - 123 -
III. XPHINGOLIPID VA GLYCOLIPID. ......................................................... - 124 -
IV. SAP.............................................................................................................. - 125 -
V. STEROL VA STEROID. ............................................................................. - 126 -
VI. SAC TO QUANG HP. .............................................................................. - 127 -
1.Chlorophyll. ................................................................................................ - 127 -
2. Caroteneoid ............................................................................................... - 128 -
3. Phycobilin. ................................................................................................. - 130 -
VII. VITAMIN TAN TRONG LIPID................................................................ - 131 -
1.Vitamin A. .................................................................................................. - 131 -
2.Vitamin D. .................................................................................................. - 132 -
3. Ubiquinone va plastoquinone. ................................................................... - 134 -
VIII. PHAN GIAI LIPID................................................................................... - 135 -
1.Phan giai lipid trung tnh............................................................................. - 135 -
2.Oxy-hoa acid beo........................................................................................ - 136 -
3. The cetone. ................................................................................................ - 142 -
4. S dung lipid d tr cho muc ch sinh tong hp. Chu trnh glyoxylate. ... - 143 -
IX. SINH TONG HP ACID BEO.................................................................... - 144 -
1. Sinh tong hp acid beo no......................................................................... - 144 -
2. Sinh tong hp acid beo khong no............................................................... - 146 -
X. SINH TONG HP TRIACYLGLYCERIN. .................................................. - 148 -
XI. SINH TONG HP GLYCEROPHOSPHOLIPID VA
GLYCEROGALACTOLIDID. .......................................................................... - 149 -
XII. SINH TONG HP SPHYINGOLIPID VA GLYCOLIPID. ...................... - 150 -
XIII. SINH TONG HP STERINE. .................................................................. - 151 -

Hoa sinh hoc - 4 -
CHNG 6. NUCLEOTIDE VA ACID NUCLEIC.............................................. - 153 -
I. NUCLEOTIDE. ............................................................................................. - 153 -
II. POLYNUCLEOTIDE ................................................................................... - 159 -
III. ADN, NHIEM SAC THE VA MAT MA DI TRUYEN. .............................. - 160 -
IV. ARN............................................................................................................. - 167 -
1.ARN thong tin (mARN). ............................................................................. - 167 -
2. ARN van chuyen (tARN)........................................................................... - 168 -
3. ARN ribosome (rARN). ............................................................................. - 171 -
V. PHAN GIAI ACID NUCLEIC. ..................................................................... - 172 -
1. Tac dung cuua exo- va endonuclease........................................................ - 172 -
2. Tac dung cua acid va kiem. ....................................................................... - 174 -
3. Phan giai nucleotide va nucleoside. .......................................................... - 174 -
4. Phan giai pentose va base nit. Cac pentose tiep tuc chuyen hoa theo con
ng chuyen hoa chung cua glucide. ........................................................... - 174 -

Hoa sinh hoc - 5 -
M AU

LOGICH PHAN T CUA VAT THE SONG
VA NHIEM VU CUA HOA SINH HOC

Hoa sinh hoc co the c xem nh hoa hoc cua cac vat the song. Moi vat the
song eu c cau tao t nhng phan t vo sinh song lai co nhng tnh chat rat ac
biet ma the gii vo sinh khong co. o la:
- Tnh phc tap va mc o to chc cao. Trong cau truc phc tap o cha vo so
cac hp chat hoa hoc vi cac kieu cau truc khac nhau. Trong khi o moi trng xung
quanh la hon hp vo trat t cua cac hp chat kha n gian;
- Moi bo phan tao thanh cua c the song (c quan, mo, te bao, cac cau truc di
te bao va cac phan t hoa hoc khac nhau) c phan cong thc hien cac chc nang xac
nh;
- Kha nang tiep nhan nang lng va nguyen lieu t moi trng va bien hoa no
e s dung cho viec xay dng va duy tr cau truc phc tap cua mnh; trong khi o cac
he thong vo sinh eu b phan huy neu chung hap thu nang lng;
- Kha nang sinh san, tc t khoi phuc mot cach chnh xac e tao ra t the he nay
en the he khac nhng ca the giong het nh mnh (neu tranh c cac yeu to gay bien
d).
La mot bo phan khong the tach ri cua t nhien, vat the song khong the khong
chu s ieu khien cua tat ca cac quy luat cua t nhien. Tuy vay, ngoai nhng quy luat
chung cua t nhien, cac phan t trong c the song con tng tac vi nhau va vi moi
trng xung quanh tren c s mot he thong cac nguyen tac ac biet ma ta co the goi
chung la logich phan t cua vat the song. o la mot he thong nhng quy luat c ban
xac nh ban chat, chc nang cua cac phan t ac biet ma ta tm thay trong c the
song va s tng tac gia chung ma nh o c the tr nen co kha nang t to chc va t
khoi phuc.
Phan ln cac thanh phan hoa hoc cua c the song la nhng hp chat hu c ma
trong o carbon ton tai dang co mc o kh cao. Nhieu phan t sinh hoc con cha
nit. Hai nguyen to nay the gii vo sinh t pho bien hn va ch ton tai dang nhng
hp chat n gian nh CO
2
, N
2
, CO
3
2-
, NO
3
-
v.v...
Cac hp chat hu c trong c the song rat a dang va phan ln la cc ky phc
tap. Tham ch c the song n gian nhat la vi khuan, v du Escherichia coli, cung a
cha ti 5000 loai hp chat hu c khac nhau, trong o co khoang 3000 loai protein va
1000 loai acid nucleic. Trong nhng c the phc tap hn ong vat va thc vat mc
o a dang con cao hn nhieu. V du, trong c the ngi co en 5 trieu loai protein,

Hoa sinh hoc - 6 -
trong o khong mot loai nao giong hoan toan vi protein cua E. coli, mac du mot so
loai co chc nang giong nhau.
Tuy nhien du cho cac phan t sinh hoc co a dang va phc tap en au, tat ca
chung eu co nguon goc rat n gian: tat ca protein eu c hnh thanh t 20 loai
aminoacid, toan bo acid nucleic t 8 loai nucleotide chu yeu. Nhng phan t vat lie u
xay dng n gian nay c chon loc trong qua trnh tien hoa e thc hien khong phai
ch mot ma nhieu chc nang e am bao nguyen tac tiet kiem toi a. Trong c the
song khong the tm thay mot hp chat nao ma khong am nhiem t nhat mot chc nang
nao o.
T nhng ieu noi tren co the rut ra mot quy luat: Tnh a dang va phc tap cua
cac phan t sinh hoc eu co coi nguon kha n gian: moi c the song eu co nguon
goc chung va c tao nen tren c s tiet kiem phan t.
Mot kha canh ac biet khac cua c the song la: bang cach nao c the song co
the tao ra va duy tr c tnh trat t va phc tap cua cau truc trong khi moi qua trnh
vat ly va hoa hoc, theo nh luat th hai cua nhiet ong hoc, eu co xu hng tien ti
cho mat trat t va hon loan, tc hng ve pha tang entropy? C the song khong the
khong tuan theo cac quy luat cua t nhien, tc chung khong the xuat hien mot cach t
phat t s hon loan. No cung khong the tao ra nang lng t cho khong co g, trai vi
nh luat bao toan nang lng. The nhng c the song co mot tnh chat ac thu quan
trong la co kha nang tiep nhan nang lng t moi trng trong nhng ieu kien nhiet
o va ap suat cu the, bien hoa nang lng o thanh dang nang lng thch hp cho ban
than chung. Nang lng hu ch ma c the song co the s dung c goi la nang lng
t do. o la phan nang lng co the tao ra cong trong ieu kien nhiet o va ap suat
khong oi. Phan nang lng ma te bao thai ra moi trng thng la dang nhiet. ieu
o gop phan lam tang entropy cua moi trng, tc lam tang tnh hon loan cua no.
Vat the song la nhng he thong h (theo cach dien at cua nhiet ong hoc), hay
nhng he thong cach ly tng oi (theo cach noi cua ieu khien hoc). Ca hai cach dien
at eu co ngha la nhng he thong nay co s lien he vi moi trng xung quanh,
trong o moi trng can cho c the song khong nhng nh nguon nang lng ma con
la nguon vat lieu xay dng. ac iem cua loai he thong nay la chung khong thiet lap
trang thai can bang vi moi trng, mac du co the cam giac rang c the ton tai trang
thai can bang v khong nhan thay co s bien oi khi quan sat chung trong mot khoang
thi gian nao o. Tren thc te th khong phai nh vay. C the song ch co the thiet lap
trang thai can bang ong vi moi trng, tc trang thai ma toc o van chuyen vat chat
va nang lng t moi trng vao he thong can bang vi toc o cua dong ngc lai.
Nh vay, te bao la mot he thong h khong can bang, mot chiec may tiep nhan nang
lng t do t moi trng e lam tang tnh trat t cua ban than, ong thi lam tang
entropy cua moi trng. May tiep nhan nang lng nay hoat ong vi hieu suat cao

Hoa sinh hoc - 7 -
hn nhieu so vi moi may moc do con ngi sang che ra. o la mat th hai cua
nguyen tac tiet kiem cua c the song. tiet kiem nang lng.
C che tiep nhan nang lng cua c the song c xay dng t nhng hp chat
hu c tng oi kem ben vng, nhay cam vi nhng ieu kien thai cc nh nhiet o
qua cao, dong ien qua manh, o pH qua lech ve pha kiem hoac acid v.v... Toan bo
he thong song, v du te bao, la mot he thong ang nhiet. V the chung khong the dung
nhiet lam nguon nang lng. Noi cach khac, te bao la nhng chiec may hoa hoc ang
nhiet. Chung ch co the thu nhan nang lng t moi trng dang hoa nang, sau o
bien hoa nang lng nay e thc hien cac cong hoa hoc trong viec tong hp cac thanh
phan cua te bao, cong tham thau trong viec van chuyen vat chat, cong c hoc trong
ong tac co c v.v...
S d te bao co the hoat ong nh chiec may hoa hoc ang nhiet la nh chung
cha mot he thong cac chat xuc tac sinh hoc gom hang ngan loai enzyme khac nhau.
o la nhng phan t protein co kha nang xuc tac mot cach ac hieu mot hoac mot so
phan ng xac nh, lam cho ieu kien ap suat va nhiet o bnh thng cua te bao cac
bien oi hoa hoc van co the xay ra vi toc o va hieu qua rat cao. Nh tnh ac hieu
cao cua enzyme ma hang loat cac phan ng khac nhau co the xay ra ong thi trong te
bao. Tnh ac hieu nay la s the hien cua mot trong nhng nguyen tac rat quan trong
cua s song nguyen tac bo sung: moi enzyme ch co the tiep nhan c chat cua mnh,
tc nhng c chat co cau tao phan t phu hp vi trung tam hoat ong cua enzyme ay.
Gia cac phan ng enzyme khac nhau trong te bao ton tai nhng moi lien he
phc tap. San pham cua phan ng nay co the la c chat cua phan ng kia. Hang loat
cac phan ng ke tuc nhau nh vay lan lt xay ra e thc hien nhng chc nang xac
nh. Nhng chuoi phan ng o con co the tao ra cac mach nhanh e gop phan hnh
thanh nen cac mang li vi nhng chc nang sinh hoc khac nhau. Toan bo nhng
mang li o ket hp vi nhau tao nen mot he thong thong nhat cac qua trnh hoa hoc
trong te bao co ten goi la trao oi chat.
Nh moi lien he chat che gia cac phan ng enzyme ma nang lng hoa hoc co
the di chuyen c trong he thong ang nhiet. Nang lng ma c the tiep nhan c
t moi trng nh moi lien he nay co the c tch luy lai bang cach phosphoryl-hoa
adenosyldiphosphate (ADP) thanh adenosyltriphosphate (ATP). Ngc lai, khi ATP b
phan giai thanh ADP se kem theo giai phong nang lng. Nh s lien he gia cac
phan ng enzyme nang lng o c s dung e tong hp cac hp chat khac nhau
hoac e thc hien mot cong nao o.
Moi lien he gia cac phan ng enzyme con la c s e tao ra cac he thong ieu
hoa trong c the song, tai o toc o cua mot phan ng ac hieu co the c ieu hoa
nh toc o cua mot phan ng khac. Nh c che ieu hoa o moi phan ng ch xay ra
theo chieu hng xac nh vi toc o xac nh u e te bao duy tr trang thai on nh
bnh thng cua mnh. Trong nhng trng hp n gian nhat s tch luy cua san pham

Hoa sinh hoc - 8 -
trung gian (chat trao oi) vi ham lng qua mc can thiet se co tac dung nh tn hieu
lam giam toc o cua chuoi phan ng tao ra chung. Cach ieu hoa nh vay c goi la
c che theo nguyen tac lien he ngc. Nhng enzyme ng au chuoi phan ng hoac
ng tai iem phan nhanh thng la nhng enzyme ieu hoa, trc tiep b c che bi
san pham cuoi cung.
Kha nang t ieu hoa cua te bao con the hien cho no t ieu khien s tong hp
he thong enzyme cua mnh. Khi te bao a nhan c t moi trng mot san pham can
thiet nao o, no se tam thi nh ch hoat ong cua he thong enzyme von can thiet e
tao ra san pham o. Ngc lai, khi tiep nhan t moi trng mot c chat can phai c
tiep tuc bien hoa, te bao se a he thong tong hp nhng enzyme can thiet cho s bien
hoa o vao hoat ong.
Cuoi cung, trong so nhng tnh chat ky dieu cua c the song, ky dieu nhat la kha
nang sinh san, tc kha nang tao ra vi mc o chnh xac hau nh tuyet oi nhng ca
the the he sau giong het the he trc. Hn the na, nhng sai sot oi khi xay ra
trong qua trnh sinh san co the lam xuat hien the he sau nhng dang ot bien khong
phai luc nao cung co hai. Ngc lai, ot bien la mot yeu to quan trong gop phan lam
cho c the song ngay cang hoan thien, la mot ong lc cua tien hoa. Kho co the tng
tng c rang mot khoi lng khong lo thong tin di truyen can e tai tao mot c the
cc ky phc tap lai co the goi gon trong nhan cua te bao trng va tinh trung be nho
dang trat t cua cac nucleotide trong phan t acid deoxyribonucleic (ADN) vi trong
lng khong qua 6.10
-12
gam. Tnh chat ky die u nay la he qua cua s phu hp ve mat
kch thc gia mat ma di truyen vi nhng bo phan tao thanh cua phan t ADN, tc
cung la he qua cua tnh bo sung ve mat cau truc. Nh nguyen tac bo sung nay ma moi
phan t ADN co the lam khuon e uc nen phan t ADN khac trong qua trnh co ten
la nhan ma (replation) hoac tao ra cac phan t acid ribonucleic thong tin (mARN)
trong qua trnh sao ma (transcription). Cung chnh nh nguyen tac nay ma mARN co
the lam khuon e uc nen cac phan t protein trong qua trnh phien ma
(translation). Ket qua la thong tin di truyen von co cau truc mot chieu dang trat t
nucleotide trong phan t ADN c bien thanh dang thong tin ba chieu ac trng
cho moi cau truc phan t va tren phan t cua vat the song. Trat t nucleotide trong
ADN quyet nh trat t aminoacid trong cac phan t protein. Mot trat t aminoacid o
cha ng nhng moi tng tac vat ly va hoa hoc phc tap, lam cho phan t protein t
ong tao ra cho mnh nhng kieu cau truc khong gian on nh va ac hieu, cho phep
chung am nhan nhng chc nang nhat nh trong he thong cac qua trnh hoat ong
song cua te bao.
Co the tom tat nhng nguyen tac a trnh bay tren ay cua logich phan t cua
vat the song mot cach ngan gon nh sau: Te bao la mot he thong ang nhiet co kha
nang t to chc, t ieu khien va t tai tao. He thong nay c hnh thanh t mot so
ln cac phan ng von lien quan mat thiet vi nhau va c thuc ay nh cac chat xuc
tac hu c do ban than te bao tao ra. Moi hoat ong cua te bao eu tuan thu mot cach

Hoa sinh hoc - 9 -
nghiem ngat nguyen tac tiet kiem toi a ve vat chat cung nh ve nang lng va thong
tin.
Logich phan t cua vat the song hoan toan khong mau thuan vi bat ky quy luat
vat ly va hoa hoc nao cung nh khong oi hoi phai phat bieu nhng quy luat mi. Tuy
nhien, ieu quan trong la cac c che cua te bao song van ch tac dung trong pham vi
cua nhng quy luat ieu khien hoat ong cua nhng may moc do con ngi tao ra,
song nhng phan ng, nhng qua trnh xay ra trong te bao song hoan thien hn nhieu
so vi nhng chiec may t ong hien ai nhat.
Con ngi ang tien gan en cho hieu biet c mot cach sau sac nguon goc va
s tien hoa cua cac phan t sinh hoc, hieu c ay u nhng phan ng enzyme ket
thanh cac qua trnh hoa hoc thong nhat trong te bao. Va khi o con ngi se hieu c
logich phan t cua vat the song xuat hien nh the nao va chng minh c nhng quy
luat cua no. Hoa sinh hoc cung vi cac lnh vc khac cua sinh hoc hien ai va vi s
ho tr cua toan hoc, vat ly hoc, hoa hoc ... ang hng ve muc tieu ay hap dan o
trong khi thc hien nhiem vu ac thu cua mnh la nghien cu nhng ac iem a c
mo ta tren ay vi cac mc o khac nhau: c the, c quan, mo, te bao, di te bao,
phan t va di phan t.


Hoa sinh hoc - 10 -

CHNG 1. AMINOACID VA PROTEIN

Protein la c s cho s hnh thanh cung nh duy tr cau truc va chc nang cua
cac vat the song nh chung co nhng ac iem ma bat ky mot hp chat hu c nao
khac cung khong the co c. o la:
- Tnh a dang vo cung cua cau truc va song song vi no la tnh ac hieu loai rat
cao;
- Tnh a dang vo cung cua cac chuyen hoa vat ly va hoa hoc;
- Kha nang tng tac noi phan t;
- Kha nang phan ng vi tac ong ben ngoai bang cach bien oi cau hnh cua
phan t theo quy luat nhat nh va khoi phuc trang thai ban au sau khi nhng tac
ong o khong con na;
- Khuynh hng tng tac vi cac hp chat hoa hoc khac e tao nen nhng phuc
he va cau truc tren phan t;
- S ton tai cua tnh chat xuc tac sinh hoc va cac hoat tnh sinh hoc khac.
Trung bnh, trong phan t protein co 50-55% C, 21-24% O, 15-18% N, 6,5-7,5%
H, 0-2,4% S, 0-2% P. Trong mot so protein con co cha Fe, Mg, I, Cu, Zn, Br, Mn, Ca
v.v... Do ham lng nit trung bnh trong protein la 16% nen e biet ham lng
protein trong mau phan tch, ngi ta thng xac nh ham lng nit roi nhan vi he
so 100/16, tc 6,25.
Protein c cau tao t 20 loai aminoacid khac nhau.
I. AMINOACID.
Aminoacid la nhng acid hu c mach beo, vong thm hoac d vong co cha t
nhat mot nhom amin (-NH
2
). Trong t nhien co khoang 150 loai aminoacid khac nhau
nhng ch co 20 loai trong so chung tham gia cau tao nen phan t protein. Nhom amin
trong 20 aminoacid nay luon gan tai nguyen t carbon -, v the chung c xep vao
nhom -aminoacid.

Hoa sinh hoc - 11 -
1. Cau tao.
Da vao cau tao va tnh chat cua goc R co the chia aminoacid thanh nhng
nhom sau ay (bang 1.1):
- Aminoacid cha goc R khong phan cc hay k nuc;
- Aminoacid cha goc R phan cc khong tch ien;
- Aminoacid cha goc R tch ien am;
- Aminoacid co goc R tch ien dng.

Bang 1.1. Cau tao cua cac aminoacid thng gap trong protein.
Ten goi
Cau thc cau tao
A. Aminoacid cha goc R khong phan cc hay k nuc
Alanine
(Ala, A)
NH
2
H
3
C -C-COOH
H
Valine
(Val)
H
3
C NH
2
CH-C-COOH
H
3
C H
Leucine
(Leu)
H
3
C NH
2
HC-CH
2
-C-COOH
H
3
C H
Isoleucine
(Ile)
NH
2
H
3
C-CH
2
-CH-CH
2
-C-COOH
CH
3
H
Proline
(Pro)
CH
2
H C
2
CH-COOH

H C
2
NH
Phenylalanine
(Phe)
NH
2
-CH
2
-C-COOH
H
Tryptophan
(Trp)
NH
2
-CH
2
-C-COOH
H
NH
Metionine
NH
2
H
3
C-S-CH
2
-CH
2
-C-COOH

Hoa sinh hoc - 12 -
(Met)
H
B. Aminoacid cha goc R phan cc khong tch ien
Glycine
(Gly, G)
NH
2
H

-C-COOH
H
Serine
(Ser, S)
NH
2
HO-CH
2
-C-COOH
H
Bang 1.1. Cau tao cua cac aminoacid thng gap trong protein (tiep theo)

Threonine
(Thr, T)
NH
2
H
3
C-CH -C-COOH
OH H
Tyrosine
(Tyr, Y)
NH
2
HO CH
2
-C-COOH
H
Cysteine
(Cys, C)
NH
2
HS-CH
2
-C-COOH
H
Asparagine
(Asn, N)
O NH
2
H
2
N-C-CH
2
-C-COOH
H
Glutamine
(Gln, Q)
O NH
2
H
2
N-C-(CH
2
)
2
-C-COOH
H
C. Aminoacid cha goc R tch ien am
Acid asparaginic
(Asp, D)
NH
2
HOOC-CH
2
-C-COOH
H
Acid glutamic
(Glu, E)
NH
2
HOOC-(CH
2
)
2
-C-COOH
H
D. Aminoacid co goc R tch ien dng
Lysine
(Lys, K)
NH
2
H
2
N-(CH
2
)
4
-C-COOH
H

Hoa sinh hoc
Khoa Sinh hoc
- 13 -

Arginine
(Arg, R)
NH NH
2
H
2
N-C- (CH
2
)
3
-C-COOH
H
Histidine
(His, H)
NH
2
-CH
2
-C-COOH
N NH H

2. Hoat tnh quang hoc.
Phan t cua moi aminoacid tr glycine eu cha t nhat mot nguyen t carbon
bat oi, do o la nhng hp chat co hoat tnh quang hoc. Chung co the ton tai cac
dang ong phan lap the D hoac L. Tuy nhien, trong t nhien hau het aminoacid eu co
dang L. C the ong thc vat noi chung ch co the hap thu c dang nay.
Cac dang ong phan D va L cua mot aminoacid co cau truc oi xng qua gng
phang. E. Fisher dien ta hai dang ong phan quang hoc nay nh sau:

Theo kieu dien at nay lien ket net am la cac lien ket tren mat phang nam
ngang hng ve pha ngi oc, con cac lien ket net nho la nhng lien ket nam pha
tren (nhom COOH) va pha di (goc R) cua mat phang nay. e thuan tien hn,
Fisher cung e xuat mot cach dien at khac goi la cong thc hnh chieu, trong o ca 4
hoa tr cua nguyen t carbon bat oi eu c bieu dien bang cac gach noi net nho
nh nhau, v tr cua cac nguyen t va nhom nguyen t c gi nguyen nh tren. V
du L-alanin c bieu dien nh sau:

GS.TS. Mai Xuan Lng
Hoa sinh hoc - 14 -
Hoat tnh quang hoc cua mot chat lien quan vi cau truc bat oi cua no, hay noi
cach khac, vi v tr cua cac nguyen t trong phan t cua chat o. Ket qua la moi dang
ong phan quang hoc co kha nang xoay mat phang cua ta sang phan cc mot goc nhat
nh sang trai (-) hoac sang phai (+). Ve mat nh lng, hoat tnh quang hoc c the
hien bang gia tr co ten goi la o quay rieng:

Gia tr nay phu thuoc nhiet o va bc song cua tia sang (thng dung tia D cua
natri vi = 5461). Trong moi day D hoac L eu ton tai nhng ai dien co hoat tnh
(+) hoac (-) vi nhng gia tr o quay rieng ac trng. Hon hp gom 50% dang D va
50% dang L cua mot aminoacid nao o c goi la rasemate. Hoat tnh quang hoc cua
hon hp o bang khong.
3. Tnh chat lng tnh.
Aminoacid co t nhat hai nhom co kha nang b ion-hoa: nhom -carboxyl vi pK
nam gia 1,7 va 3,0 va nhom -amin vi pK vao khoang 10. Trong dung dch co pH
gia 4 va 7 aminoacid ton tai dang ion lng tnh (zwisterion) khi ca hai nhom tren
eu b ion hoa:
R CH COOH H
+
+ R CH COO
-
pH = 1,7 2,8
NH
3
+
NH
3
+
R CH COO
-
H
+
+ R CH COO
-
pH = 9 10,7
NH
3
+
NH
2
Ngoai ra, mot so aminoacid con cha cac nhom ion-hoa khac:
- Cac nhom - va -carboxyl cua acid asparaginic va acid glutamic:
R COOH H
+
+ COO
-
pH = 4,3
- Nhom -imidazol cua histidine:
NH
+
N
H
+
+ pH = 6,1
NH NH
- Nhom -amin cua lysine:
R NH
3
+
H
+
+ NH
2
pH = 10,5
- Nhom -sulfhydril cua cysteine:
R - SH H
+
+ R S
-
pH = 8,1 8,3
- Nhom -guanidine cua arginine:

Hoa sinh hoc - 15 -
H
2
N C NH R H
+
+ H
2
N C NH R pH = 12,5
NH
2
+
NH
Hnh 1.1 trnh bay ng cong chuan o mo ta s ion-hoa ke tiep nhau cua
lysine (aminoacid co tnh base) va acid asparaginic (aminoacid co tnh acid).
Qua trnh phan ly cua cac nhom ion-hoa trong phan t cua hai chat nay c
minh hoa cac loat phan ng (1) va (2) :

Hnh 1.1. ng cong chuan o cua lysine va acid asparaginic.

(1) COOH OH
-
COO
-
OH
-
COO
-
OH
-
COO
-
CHNH
3
+
H
+
CHNH
3
+
H
+
CHNH
2
H
+
CHNH
2
(CH
2
)
3
pK1=2,18 CH
2
)
3

pK
2
=8,95 (CH
2
)
3

pK
3
=10,53 CH
2
)
3
CH
2
NH
3
+
CH
2
NH
3
+
CH
2
NH
3
+
CH
2
NH
2
(2) COOH OH
-
COO
-
OH
-
COO
-
OH
-
COO
-
CHNH
3
+
H
+
CHNH
3
+
H
+
CHNH
3
+

H
+
CHNH
2
CH
2
pK1=2,01 CH
2
pK
2
=3,80 CH
2
pK
3
=9,93 CH
2
COOH

COOH COO
-
COO
-

pH sinh ly ca hai nhom NH
2
cua lysine cung nh nhom COOH cua no eu
b ion-hoa, tao ra mot ien tch dng yeu trong phan t. Cung gia tr pH nay acid
asparaginic mang ien tch am yeu do s phan ly cua cac nhom carboxyl cua no.

Hoa sinh hoc - 16 -
Cac gia tr pK cua cac aminoacid khac nhau c gii thieu trong bang 1.2.
Can nh rang pK la gia tr am cua loga thap phan cua hang so phan ly K, trong
o:
[A
-
]
K = [H
+
] x (HA la aminoacid trang thai khong phan ly)
[HA]
Theo phng trnh Henderson-Hasselbalch, pH, tc gia tr am cua loga thap
phan
[A
+
]
cua [H
+
], bang pK + log .
[HA]

Bang 1.2. Gia tr pK cua mot so aminoacid.
Aminoacid PK
1
PK
2
PK
3
Aminoacid PK
1
PK
2
PK
3
Gly 2,35 9,78 - Pro 2,20 10,60 -
Ala 2,34 9,87 - HyPro* 1,92 9,73 -
Ser 2,21 9,15 - Glu 2,19 4,28 9,66
Cys 1,96 8,18 10,28 Asp 2,09 3,87 8,82
Met 2,28 9,21 - His 1,77 6,10 8,17
Val 2,32 9,62 - Lys 2,18 8,95 10.53
Leu 2,36 9,60 - Arg 2,09 9,04 12,48
Ile 2,36 9,68 - Thr 2,63 10,40 -
Tyr 2,60 9,10 10,10 Gln 2,17 9,13 -
Phe 2,58 9,24 - Asn 2,02 8,80 -
Trp 2,38 9,39 * - HyPro - Hydroxyproline
Gia tr pH ma tai o phan t aminoacid (hoac mot chat ien phan lng tnh)
mang so ien tch am va dng nh nhau c goi la iem ang ien va ky hieu la
pH
i
. Tai gia tr nay aminoacid khong di chuyen trong ien trng mot chieu. Moi
aminoacid co iem ang ien ac trng.
Thong thng, gia tr cua pH
i
cua mot aminoacid la gia tr trung bnh cong cua
cac gia tr pK cua no.
4. Cac phan ng hoa hoc ac trng.
Tnh chat hoa hoc ac trng cua aminoacid bao gom cac phan ng lien quan vi
nhom -carboxyl, -amin va cac nhom chc khac trong thanh phan cua goc R.
a/ Cac phan ng cua nhom -carboxyl:
- Phan ng vi kiem e tao ra muoi tng ng:

Hoa sinh hoc - 17 -
R CH COO
-
+ NaOH R CH COONa + H
2
O
NH
2
NH
2
- Phan ng vi ru e tao ester:
R CH COOH + CH
3
- CH
2
OH R CH C O CH
2
-CH
3
+ H
2
O
NH
2
N O
Nhnbg ester nay co the ngng tu khi un nong e tao ra pan pham mau
dicetopyperasine tng ng:
CHR
2 R CH COOR O = C NH + 2R -OH
NH
2
HN CO
CHR
Di tac dung cua cac tac nhan kh manh, v du borhydrite natri, aminoacid b
kh thanh ru amine tng ng:
NaBH
4
R CH COOH + CH
3
- CH
2
OH R CH CH
2
OH
NH
2
NH
2

- Ket hp vi ammoniac e tao ra amide:
NaBH
4
R CH COOH + NH
3
R CH C - NH
2
NH
2
NH
2
O
Hai phan ng nay cung thng c s dung e xac nh trat t aminoacid trong
chuoi polypeptide t au tan cung cha nhom COOH t do (C-tan cung).
b/ Cac phan ng cua nhom -amine.
- Phan ng vi HNO
2
tao nen oxyacid tng ng, nit t do va nc:
R CH COOH + HNO
2
R CH COOH + N
2
+ H
2
O
NH
2
OH
Phan ng vi aldehyde, v du pormaldehyde, e tao ra cac hp chat rat linh ong
co ten chung la hp chat schiff:
R CH COOH + CH
2
O

R CH COOH + H
2
O
NH
2
N=CH
2

Hai phan ng nay thng c dung e nh lng aminoacid t do.
- Phan ng vi ninhydrin:
Khi un nong vi ninhydrin a so aminoacid b oxyhoa va phan giai thanh
anhydride tng ng, CO
2
va NH
3
.

Hoa sinh hoc - 18 -

Trong nhng ieu kien nhat nh NH
3
ngng tu vi dicetohydrinden va vi phan
t ninhydrin th hai e tao thanh san pham mam tm o hay lam o:

Phan ng nay rat nhay va c dung e nh lng aminoacid bang phng
phap sac ky va ien di tren giay.
Proline va oxyproline tao ra vi ninhydrin cac san pham mau vang.
- Phan ng Sanger:

Phan ng nay c Sanger dung e xac nh trat t aminoacid cua chuoi
polypeptide t au N-tan cung, tc au tan cung cha nhom NH
2
t do.
Phan ng vi 1-dimethylaminophtalin--sulfonylchloride (goi tat la dansyl-
chloride):

- Phan ng Edman:
-Aminoacid phan ng vi phenylisothiocianate e tao ra cac dan xuat
phenylthio-hydatoyl tng ng:

Hoa sinh hoc - 19 -

Hai phan ng cuoi cungc s dung vi muc ch nh phan ng Sanger.
c/ Cac phan ng cua goc R.
Goc R cua cac aminoacid khac nhau cha cac nhom hoat ong khac nhau nh a
gii thieu muc (1). Cac phan ng vi s tham gia cua chung rat a dang va la yeu to
quan trong gop phan xac nh ac iem cau truc va hoat tnh sinh hoc cua protein.
Chung se c xem xet ti sau trong cac muc IV va VI.
II. PEPTIDE.
Peptide la nhng hp chat c hnh thanh t t nhat hai phan t aminoacid,
trong o nhom -amine cua aminoacid nay ket hp vi nhom -carboxyl cua
aminoacid kia bang lien ket -CO-NH- (lien ket peptide):
Tuy thuoc so goc aminoacid trong phan t, ngi ta phan biet di-, tri-,
tetrapeptide v.v... Nhng peptide cha t hai en khoang 10 goc aminoacid c goi
chung la oligopeptide. Neu so goc aminoacid at en hang tram, ta co polypeptide
c s cau truc cua protein.
e goi ten cac peptide phan t nho, ngi ta ghep lan lt t au N-tan cung ten
cua cac goc aminoacid, trong o uoi -ine c oi thanh -yl; rieng ten cua
aminoacid C-tan cung c gi nguyen. V du glycylalanin, glycylalanylserin v.v...
Trong t nhien a phat hien c nhieu peptide khac nhau co vai tro sinh hoc rat
quan trong nh glutation, oxytosin,vasopresin...
Glutation (-glutamylcysteylglycine) la mot tripeptide co cau tao nh sau:
HOOC CH CH
2
CH
2
CO NH CH CO NH CH
2
C00H
NH
2
CH
2
SH
Goc acid glutamic Goc cysteine Goc glycine
Glutation co vai tro quan trong trong ho hap. Tr dang kh tren ay no co the
chuyen hoa thuan nghch thanh dang oxy hoa:

Hoa sinh hoc - 20 -
HOOC CH CH
2
CH
2
CO NH CH CO NH CH
2
C00H
NH
2
CH
2
S
S
NH
2
CH
2
HOOC CH CH
2
CH
2
CO NH CH CO NH CH
2
C00H

Nh s chuyen hoa thuan nghch nay ma glutation co the tham gia trong qua
trnh van chuyen ien t trong te bao. Oxytosin va vasopresin la cac hormone cua
thuy sau tuyen yen. Chung c hnh thanh t 9 goc aminoacid.
Vasopresin co tac dung lam tang
huyet ap, c che s hnh thanh nc
tieu. Oxytosin anh hng en s co
bop cua da con va cac c trn khac

Hoa sinh hoc - 21 -
III. TNH CHAT CUA LIEN KET PEPTIDE.
Lien ket C-N trong lien ket
peptide khong the xoay t do va mang
tnh chat trung gian gia lien ket oi va
lien ket n. Khoang cach gia C va N
trong lien ket peptide bang 0,13nm, tc
ngan hn lien ket bnh thng gia
nguyen t carbon - va nguyen t nit
ke can (0,15nm va dai hn lien ket oi
thc thu gia C va N (0,125nm).
Bon nguyen t cua lien ket
peptide va hai nguyen t carbon -
luon nam tren cung mot mat phang,
trong o oxy cua nhom carbonyl va
hydro cua nhom amin nam v tr trans
oi vi nhau.

Hnh 1.2. Cau tao cua lien ket peptide.
Hnh I.2. Cau tao cua lien ket peptide.

Goc R va nguyen t hydro nh vi C- nam hai pha oi dien cua mat phang
va tao vi mat phang nay mot goc 109
o
28. Khac vi lien ket gia C va N trong lien
ket peptide, lien ket gia C- va N trong chuoi peptide co the xoay t do va quyet
nh dang cau truc cua phan t protein (hnh 1.2).
Cung do tnh chat trung gian nay nen chuoi peptide co the ton tai ca hai dang
cetone va enol (tnh chat ho bien):
...-NH-CH-C-NH-CH-C-NH-... ...=N-CH-C=N-CH-C=N-...
R O R O R OH R OH
Nh tnh ho bien nay ma kha nang phan ng cua phan t protein tang len ang
ke.
IV. CAC LIEN KE T TH CAP TRONG PHAN T PROTEIN.
Ngoai lien ket peptide co nhiem vu noi cac aminoacid vi nhau e tao nen chuoi
polypeptide, trong phan t protein con xuat hieh nhieu kieu lien ket th cap gop phan
lam on nh phan t va quy nh dang cau truc cua phan t. Nhng lien ket nay hnh

Hoa sinh hoc - 22 -
thanh c la do s ton tai cua cac nhom chc rat a dang trong cac goc R vi nhng
kha nang phan ng va tng tac khac nhau (hnh 1.3):
Lien ket ion gia cac nhom tch ien trai dau;
Lien ket disulfide gia cac nhom SH cua cysteine thuoc cung mot chuoi
polypeptide hoac gia cac chuoi polypeptide khac nhau;
Lien ket ester gia nhom hyroxyl cua serine hay threonine vi nhom - va -
cua cac aminoacid dicarboxylic;
- Lien ket hydro gia cac nhom C=O va N-H thuoc cac lien ket peptide hoac
gia H va O trang thai lien ket ong hoa tr thuoc cac nhom chc trong mot so goc R
(v du gia nhom -COOH cua goc acid glutamic va nhom -amin cua goc lysine).
- Lien ket k nc gia cac nhom khong phan cc (co tnh k nc) chi chung b
dung moi (nc) xo ay. Trong trng hp o chung co xu hng tap hp lai va gan
bo vi nhau. Cung nh lien ket hydro, lien ket k nc co nang lng khong ln. Tuy
vay, do ton tai vi so lng nhieu nen vai tro cua hai kieu lien ket nay, ac biet la lien
ket hydro, khong kem quan trong so vi cac kieu lien ket th cap khac trong viec xac
nh cau truc, tnh chat va chc nang cua protein.

Hnh 1.3. Cac kieu lien ket th cap trong phan t protein: (1) Lien ket hydro; (2)
Lien ket k nc; (3) Lien ket ion (4) Lien ket disulfide; (5) Lien ket
ester.

Trong viec lam on nh cau truc cua phan t protein con co vai tro cua lc
Vandervaals, tc lc hap dan gia cac nguyen t xuat hien do s thang giang ien the
von c cam ng khi khoang cach gia chung va u e lam xuat hien lc hap dan
tnh ien gia cac ien t tch ien am cua nguyen t nay vi nhan tch ien dng
cua nguyen t khac. Do nhan b bao boc bi lp ien t nen lc Vandervaals rat yeu
(1-2 Kcal/mol). Trong ieu kien nhiet o sinh ly lc hap dan Vandervaals se co hieu
lc nhat neu nhieu nguyen t cua mot phan t tng tac mot cach ong thi vi nhieu
nguyen t cua phan t khac.


Hoa sinh hoc - 23 -
V. CAU TRUC CU A PROTEIN.
1.Cau truc bac mot.
Ngi ta goi cau truc bac mot cua protein la thanh phan va trat t sap xep cua
cac goc aminoacid trong chuoi polypeptide. Tnh a dang cua protein c the hien
chu yeu tnh a dang trong cau truc bac mot cua chung, v moi loai protein co cau
truc bac mot hoan toan xac nh va ac trng. Mot s nham lan nho trong cau truc bac
mot lien quan en mot trong so hang tram goc aminoacid cung u e lam cho tnh chat
cua protein bien oi ang ke va gay ra nhng benh phan t rat tram trong. V du ien
hnh cho loai benh phan t nay la benh thieu mau hong cau liem. Hemoglobin cua
ngi mac benh nay (HbS) ch khac vi hemoglobin cua ngi khoe (HbA) cho goc
acide glutamic v tr th 6 cua mot trong hai chuoi polypeptide cua hemoglobin
(chuoi -) a b thay the bi goc valine:
HbAH
2
N-Val-His-Leu-Thr-Pro-Glu-Glu-Lys-....
HbS:H
2
N-Val-His-Leu-Thr-Pro-Val-Glu-Lys-....
S nham lan nay a lam cho tnh chat ly hoa cua hemoglobin bien oi. T
dang hnh cau no chuyen thanh dang li liem va khong con kha nang van chuyen oxy
na.
Xac nh cau truc bac mot cua protein c thc hien chu yeu tren c s cac
phan ng Sanger, Dansyl, Edman va mot so phan ng khac ket hp vi phng phap
ien di, sac ky, thuy phan bang enzyme v.v... ac biet, phan ng Edman la c s hoat
ong cua may phan tch aminoacid t ong, v sau khi tao ra phenylhydantoyl chuoi
peptide va b rut ngan mot goc aminoacid t au N-tan cung lai tiep tuc vong phan
ng khac vi phenylisothiocianate.
2. Cau truc bac hai.
Danh t cau truc bac hai c dung e ch cac trang thai cau truc xoan - va
duoi thang - cua chuoi polypeptide (hnh 1.4).
Cau truc xoan - la kieu sap xep trong khong gian quan trong va thng gap
nhat cua cac phan t protein hnh cau. No cung ac trng cho -keratin cua toc, long
cu, mong, sng (protein si). Chuoi polypeptide cuon xung quanh mot truc tng
tng theo chieu quay phai (chieu kim ong ho neu nhn t au N-tan cung). Moi
vong xoan gom 3,6 goc aminoacid. Nh vay, c 18 goc aminoacid tao ra 5 vong xoan
tron ven. Moi bc cua mach xoan dai 0,54nm, do o chieu dai cua moi aminoacid
tren truc la 0,54 : 3,6 = 0,15nm. Cau truc nay hnh thanh c la do s sap xep tren
mat phang mot cach co nh cua lien ket peptide va kha nang xoay t do cua C ke
can. Hai mat phang co chung nguyen t C giao nhau vi goc 108
o
. S ton tai cua vo
so cac lien ket hydro trong chuoi polypeptide lam cho cau hnh cua phan t c on

Hoa sinh hoc - 24 -
nh. Cau truc on nh nhat cua phan t protein co c khi so lng lien ket hydro
ln nhat.
Cac goc R toa ra t chu vi cua ong truc tng tng. Noi chung, cac goc
aminoacid k nc xoay vao pha trong, con nhng goc a nc hng ra ngoai. Trong
cac phan t protein hnh cau khu vc xoan ch chiem mot phan nhat nh (v du 75%
myoglobin), phu thuoc pH cua moi trng va thanh phan aminoacid. Nhng khu vc
cha isoleucine, proline va glycine khong co cau truc xoan .

Hnh 1.4. Cac kieu cau truc bac hai cua protein.
Cau truc duoi ton tai trong fibroin cua t va -keratin. Theo kieu cau truc nay
cac chuoi polypeptide trang thai hau nh duoi thang nam song song nhau va ngc
chieu nhau. Trai vi cau truc , trong cau truc cac lien ket hydro hnh thanh gia
cac chuoi polypeptide khac nhau. Cac goc R hng thang goc vi mat phang lien ket
peptide ve pha tren va pha di. Phoi hp mot so day nh vay se tao nen cau truc
dang to ong cua protein.
Ngoai hai kieu cau truc tren ay con co mot kieu cau truc bac hai khac ac trng
cho protein si colagen. Loai protein nay cha 1/3 glycine va proline nen khong the
tao nen cau truc . Ket qua phan tch cau truc bang tia Rng-ghen cho thay trong si
colagen c 3 chuoi polypeptide cuon lai vi nhau tao ra mot dang n v co ten goi la
tropocolagen. Cac n v tropocolagen nay sap xep trong si colagen cua gan theo
tng bac lech nhau khoang 60-70nm.

Hoa sinh hoc - 25 -
3. Cau truc bac ba.
Cau truc bac ba la cau hnh khong gian cua chuoi polypeptide. No c xac nh
bi cau truc bac mot va bac hai. Cau truc bac ba hnh thanh mot cach t phat phu
thuoc vao kch thc, hnh dang va tnh phan cc cua cac goc aminoacid. Nhng goc
nay tng tac vi nhau va vi cac phan t dung moi va bang cach o lam suy yeu kha
nang xoay t do cua cac lien ket trong chuoi polypeptide. S tng tac o c thc
hien bang cac kieu lien ket disulfide, ester, hydro va tng tac k nc.
Cac chuoi polypeptide cua protein hnh cau c cau tao thanh khoi mot cach
ran chac. V du phan t myoglobin (gom mot chuoi polypeptide duy nhat vi trong
lng phan t 16.700, cha 153 goc aminoacid) c bo chat en mc trong long no
ch co the cha 4 phan t nc. Tat ca cac goc R co tnh phan cc c sap xep mat
ngoai dang hydrate-hoa con hau het cac goc R khong phan cc co tnh k nc nam
trong long phan t e tranh tiep xuc vi nc. Nhng goc aminoacid khong co kha
nang tao cau truc xoan nam tai nhng ni chuoi polypeptide b gap khuc.
Tnh chat sinh hoc cua protein phu thuoc khong nhng vao cau truc bac mot, bac
hai ma ca vao cau truc bac ba cua chung. Cac yeu to pha v cau truc bac ba eu lam
mat hoat tnh sinh hoc cua protein.
4. Cau truc bac bon.
Phan t cua a so protein vi trong lng phan t tren 50.000 thng c c
cau tao t hai chuoi polypeptide tr le n. Nhng chuoi polypeptide nay c gan vi
nhau bang cac loai lien ket yeu nh lien ket hydro, lien ket k nc hoac cac loai lien
ket manh hn nh lien ket ester, lien ket disulfide v.v... Moi chuoi polypeptide vi
cau truc bac mot, bac hai va bac ba xac nh tao thanh mot phan di n v. Nhng
phan di n v o ket hp vi nhau e tao nen phan t protein hoan chnh. Cac sap
xep ac trng trong khong gian cua cac phan di n v trong moi phan t protein
hoan chnh loai nay c goi la cau truc bac bon cua protein o.
V du ien hnh cho nhng protein co cau truc bac bon la hemoglobin
(M=68.000) c cau tao bi 4 chuoi polypeptide - 2 chuoi va 2 chuoi . Chuoi
cha 141 goc aminoacid, con chuoi - 146 goc. Moi chuoi gan vi mot phan t hem.
Ca 4 phan di n v sap xep tng ho nhau theo quy luat hoan toan xac nh, lam
cho phan t hemoglobin co dang gan nh hnh cau vi kch thc 50 x 50 x 64 .
Trong nhieu trng hp cac phan di n v co the hnh thanh t mot so chuoi
polypeptide. Khi o moi chuoi polypeptide c goi la protomer, con phan di n v
c goi la oligomer. Nhng oligomer nay ket hp vi nhau thanh mot phan t protein
co cau truc bac bon hoan chnh. o la trng hp cua glutamate dehydrogenase cua
gan bo. Enzyme nay (M=2,2 x 10
6
) cau tao bi 8 oligomer vi M=280.000. Moi
oligomer nay lai c hnh thanh t mot so chuoi polypeptide vi trong lng phan t
khoang 50.000.
Ban than hemoglobin cung co the phan ly thanh 2 oligomer, moi oligomer cha
mot chuoi va mot chuoi .

Hoa sinh hoc - 26 -
Cac phan t protein co cau truc bac bon trong nhng ieu kien nhat nh phan ly
thanh cac phan di n v; trong nhng ieu kien khac nhng phan di n v nay
lai ket hp vi nhau thanh nhng phan t ban au. Qua trnh phan ly va ket hp nay
kem theo s bien oi tnh chat sinh hoc cua protein. Hoat tnh sinh hoc cung phu thuoc
vao cac kieu to hp khac nhau cua cac phan di n v. Moi lien quan nay gia cau
truc bac bon va tnh chat cua protein la c s cua nhieu qua trnh ieu hoa trong te
bao. Nhng enzyme then chot mang chc nang ieu hoa cac qua trnh trao oi chat
eu la nhng enzyme co cau truc bac bon.
S ton tai protein co cau truc bac bon con la yeu to gop phan khac phuc tac hai
cua nhng s nham lan trong qua trnh sinh tong hp protein. o cung la mot phng
tien e tiet kiem ADN va ARN thong tin.
VI. TNH CHAT CUA PROTEIN.
1. Tnh chat lng tnh.
Trong dung dch, cung nh aminoacid, protein ton tai dang ion lng cc. Tuy
nhien, khac vi aminoacid, tnh chat lng tnh cua protein c quyet nh bi cac
goc R co kha nang ion-hoa. Cac nhom -carboxyl va -amin tan cung do chiem ty le
rat t trong phan t protein nen anh hng khong ang ke en tnh chat nay. Moi
protein co mot iem ang ien ac trng. Tai iem ang ien protein rat de b ket tua.
Tnh chat lng tnh c s dung e tach va tinh che protein tren c s phng phap
ien di.
2. Hoat tnh quang hoc.
Protein la nhng hp chat co hoat tnh quang hoc do trong phan t cha nhieu
nguyen t carbon bat oi. Tuy nhien, o quay tong so cua chuoi polypeptide khong
phai la phep cong n gian o quay cua toan bo aminoacid trong thanh phan cua no.
Protein thng co o quay phai ln hn. Mc quay phai tha toi a c ghi nhan
khi chuoi polypeptide co dang xoan . Khi chuoi polypeptide trang thai hon on th
khong nhan thay co s chenh lech nay. V vay, tren c s hoat tnh quang hoc co the
xac nh ty le khu vc xoan trong phan t protein va nghien cu cac yeu to anh
hng en s hnh thanh va pha v cau truc o.
3. Tnh hydrate-hoa.
Protein lien ket chat che vi nc trong c the nh phan t cua chung co cha
cac nhom a nc nh COOH, -OH, -NH
2
v.v... Nhng nhom nay loi cuon cac phan
t nc von co tnh phan cc ve pha mnh. Nh o hnh thanh mot lp nc xung
quanh phan t protein. Nh vay dung dnh protein kha ben vng va kho b ket tua.
Hien tng nay c goi la s hydrate hoa. Chnh nh lp nc hydrate-hoa bao boc
xung quanh nen mac du tai iem ang ien protein de b ket tua nhng noi chung t
ban than no se khong b ket tua. Neu loai tr lp nc nay cac phan t protein se co

Hoa sinh hoc - 27 -
ieu kien tiep can nhau, phoi hp lai thanh nhng hat ln e cuoi cung vt khoi gii
han kch thc cua cac hat keo va do o b ket tua.
Viec rut nc co the thc hien nh muoi (diem tch) hoac cac dung moi hu c
co tnh hut nc manh nh ethanol, acetone v.v... Neu viec rut nc c thc hien
trong ieu kien em du (v du nhiet o thap), th tua protein van gi c kha nang
hoa tan tr lai cung vi ay u cac tnh chat nguyen thuy cua no.
Dung dch protein trong nc la mot dang dung dch keo (sol). Trong nhng ieu
kien nhat nh sol b mat nc va bien thanh dang keo ac (gel). Gel khi tiep xuc vi
nc se hut mot lng nc rat ln va trng phong. Kha nang trng phong nay thap
nhat tai iem ang ien. S trng phong cua hat khi ngam nc va nay mam, s
trng phong cua bot nhao v.v...eu lien quan chat che vi s trng phong cua gel
protein. Khi b bien tnh, kha nang hut nc va trng phong cua protein b giam hoac
mat hoan toan. Mat khac, trang thai gel protein kho b bien tnh hn trang thai sol.
4. S bien tnh cua protein.
Phan t protein co the b bien tnh bi nhiet o cao, pH thai cc, tia X, tia cc
tm, tia phong xa, ap lc cao, cac tac ong c hoc (v du lac manh dung dch), tac ong
cua mot so hoa chat v.v...
S bien tnh xay ra do cau truc nguyen thuy b pha v, chu yeu v t cac lien
ket th cap nh lien ket hydro, lien ket disulfide v.v... con cau truc bac mot van con
gi nguyen ven. Bien tnh keo theo hien tng mat kha nang hoa tan (ket tua), bien
oi hoat tnh quang hoc, mat cac tnh chat sinh hoc, xuat hien cac yeu to khang
nguyen mi va tr nen de b phan giai hn bi cac emzyme tieu hoa.

Hnh 1.5. S bien tnh thuan nghch cua protein. 1- bien tnh;
2- khoi phuc trang thai nguyen thuy; gach noi am lien ket disulfide.
Khi s bien tnh xay ra cha hoan toan va neu sau o loai bo dan tac nhan gay
bien tnh (v du giam nhiet o t t), trang thai nguyen thuy cua protein co the c
khoi phuc. Kieu bien tnh nay c goi la bien tnh thuan nghch (hnh 1.5).

Hoa sinh hoc - 28 -
Khi protein a bien tnh hoan toan va sau sac, cung nh khi giam nhiet o ot
ngot, protein bien tnh kho hoac khong the khoi phuc trang thai nguyen thuy (bien tnh
khong thuan nghch).
5. Cac phan ng mau ac trng.
Tuy thuoc vao thanh phan aminoacid, protein co the cho cac phan ng mau khac
nhau.
a/ Phan ng biure. Khi cho kiem va sulphate ong va dung dch protein se tao
san pham mau xanh tm hoac tm o tng t nh khi nhng chat nay tac dung vi
biure (H
2
N-CO-NH-CO-NH
2
). Phan ng nay, v the, chng minh s ton tai cua lien
ket peptide trong phan t protein.
b/ Phan ng xanthoprotein. Di tac dung cua acid nitric ac mot so protein cho
san pham mau vang. Phan ng ac trng cho nhng protein co cha cac aminoacid co
nhan benzol (tyrosine, phenylalanine, tryptophan).
c/ Phan ng Milon. Khi un protein vi thuoc th Milon (hon hp nitrate va
nitrite trong acid nitric ac) se xuat hien san pham mau hong hoac o tham. Phan ng
lien quan vi s ton tai cua nhom phenol cua tyrosine.
d/ Phan ng Adamkjevich. Nhng protein co cha tryptophan do s ton tai cua
nhan indol nen khi tac dung vi acid glyoxylic trong acid sulfusuric se cho san pham
mau xanh tm.
e/ Phan ng Sacagusa. Mot so protein khi x ly trc bang hypoxhloride natri va
sau o bang -naphtol se cho mau o tham. Phan ng cho biet trong phan t co nhom
guanidine cua arginine.
6. Hoat tnh va chc nang sinh hoc cua protein.
Hoat tnh sinh hoc cua protein the hien nhieu kha canh khac nhau. Ben canh
nhng protein cau truc vi chc nang chu yeu la gop phan tao nen cac c quan t cua
te bao, con co nhng protein enzyme lam nhiem vu xuc tac cac phan ng hoa hoc cua
qua trnh trao oi chat, Nhieu protein khac lam nhiem vu van chuyen (hemoglobin,
myoglobin), c ong (myosin, actin), bao ve (protein khang the, fibrinogen, trombin),
hormone ( insulin, hormone sinh trng), d tr chat dinh dng (ovoalbumin, casein,
glyadin, zein). Mot so protein la oc to rat manh (noc ran, risin, oc to cua mot so
nam va vi khuan). Ngoai ra, nh hoat tnh sinh hoc cua mnh, protein con thc hien
nhieu chc nang khac trong te bao va c the song. Sau khi b bien tnh, protein khong
con co kha nang am nhan bat ky cha nang sinh hoc nao.
Hoat tnh sinh hoc cua protein , cung nh tnh chat vat ly va hoa hoc cua chung,
phu thuoc vao thanh phan aminoacid cua chuoi polypeptide. Tnh a dang cua cac goc
aminoacid trong phan t protein cho phep hnh thanh cac kieu lien ket th cap khac
nhau trong noi bo phan t protein, gia cac phan t protein vi nhau va gia cac phan

Hoa sinh hoc - 29 -
t protein vi cac phan t thuoc cac nhom hp chat khac nham xay dng cac to hp
tren phan t can thiet e tao nen cac c quan khac nhau va quy nh tnh chat vat ly,
hoa hoc cung chc nang sinh hoc cua nhng to hp o.
Nhieu yeu to khac nhau cua moi trng nh nhiet o, pH v.v... co tac dung lam
thay oi moi tng quan gia cac goc aminoacid trong phan t protein. S thay oi o
dan en hang loat nhng bien oi ve mat tnh chat vat ly, hoa hoc, sinh hoc cua
protein. Mot trong nhng v du ve s bien oi o la s bien tnh.
VII. PHAN LOAI PROTEIN.
Do tnh a dang ve mat cau truc va chc nang cua protein nen oi vi nhieu loai
protein khong the thc hien phan loai mot cach thoa ang. Noi chung, co hai cach
phan loai protein khac nhau. Cach th nhat la da vao hnh dang, tnh tan, chc nang
va thanh phan hoa hoc e chia protein thanh nhieu nhom. Da vao thanh phan hoa
hoc, chia protein thanh hai nhom ln: protein n gian va protein phc tap.
Protein n gian da vao tnh tan c chia thanh cac nhom nho sau ay:
Albumin: tan trong nc, b ket tua nong o muoi kha cao (70-100). Nhom
protein nay pho bien c the ong vat va thc vat. Tinh the albumin long trang trng
va albumin huyet thanh c s dung rong rai trong nhieu lnh vc.
Globulin: khong tan hoac tan rat t trong n c, tan trong dung dch loang cua
cac muoi trung tnh. Cac protein thuoc nhom nay thng b ket tua tromg dung dch
(NH
4
)
2
SO
4
ban bao hoa. Globulin co trong huyet thanh mau, long trang trng. thc
vat globulin co trong la va trong hat cua cac cay ho au. hat hoa thao tap trung chu
yea trong tang aloron cua hat.
Prolamin: khong tan trong nc va dung dch muoi loang nhng tan trong
ethanol ho85c isopropanol 70-80%. Prolamin hau nh ch co trong noi nhu chc tinh
bot cua hat hoa thao, v du gliadin cua hat lua my, hordein cua ai mach, zein cua
ngo.
Glutelin: Ch tan trong dung dch kiem hoac acid loang. Glutelin co trong noi
nhu cua hat hoa thao va hat cua mot so cay khac, v du glutelin cua hat lua my,
oyzenin cua hat lua.
Prolamin va glutelin la cac protein d tr ien hnh cua hat hoa thao. Chung ket
hp vi cac thanh phan khac trong noi nhu cua hat tao thanh mot phc hp co trong
lng phan t rat ln, goi la gluten vi cau truc khong gian vo cung phc tap.
Histon: Protein co tnh base do cha nhieu cac aminoacid co tnh base nh
lysine va arginine., de tan trong nc, khong tan trong dung dch ammoniac loang. Se
trnh bay ky hn nhom protein nay trong chng acid nucleic.
Ngay nay ngi ta co xu hng chia protein thanh hai nhom ln: protein hnh
cau va protein si. Protein hnh cau co dang hnh cau hoac hnh elp. Hnh dang nay
co the xuat hien t cau hnh cua mot hoac mot so chuoi polypeptide. Trong mot so
trng hp (thng goi la protein n gian) nhng chuoi polypeptide nay khong ket

Hoa sinh hoc - 30 -
hp vi bat ky thanh phan nao khac. Trong mot so trng hp khac chuoi polypeptide
ket hp vi mot chat ac biet kho ng co ban chat aminoacid goi la nhom them, hay
nhom prostetic. Nhng protein loai nay thng c goi la protein phc tap.
Protein phc tap thng c mo ta theo nhom nhom them cua chung, v du,
hemoglobin thuoc nhom hemeprotein, co nhom nhom them la heme; con cac protein
cha lipid, glucid, kim loai, acid nucleic c xep tng ng vao cac nhom
lipoprotein, glycoprotein, metaloprotein hoac nucleoprotein.
Nucleoprotein: Nhom nhom them la acid nucleic, apoprotein la cac protein co
tnh base, v vay chung ket hp vi nhau kha chat. Muon tach rieng chung phai dung
dung dch muoi hoac acid loang. Nucleoprotein tap trung trong nhan te bao va
ribosome.
Nucleoprotein trong tinh dch ca do acid nucleic ket hp vi protamin, mot
polypeptide co trong lng phan t gan bang 5.000 dalton, co tnh base do cha nhieu
arginine.
Chromoprotein: la protein phc tap cha nhom nhom them co mau. Tuy theo
ac tnh cua nhom prosteic ma cac chromoprotein khac nhau co cac mau khac nhau, v
du hem (porphyrin cha sat) co mau o la nhom nhom them cua hemoglobin,
mioglobin, cytochrome C, catalase; riboflavine co mau vang, la nhom nhom them cua
cac flavoprotein.
Cac chromoprotein co hoat tnh sinh hoc cao, tham gia trong nhieu qua trnh
song quan trong nh ho hap, cac phan ng oxy hoa kh cua qua trnh thu nhan anh
sang (rodopsin)
Lipoprotein: Nhom them la lipid. Lipoprotein ong vai tro quan trong trong
qua trnh van chuyen lipid trong c the. Lipid khong tan trong nc, nhng sau khi ket
hp vi protein, phan lipid k nc cuon vao trong, phan apoprotein a nc lam
thanh lp vo boc ben ngoai nen co the c van chuyen trong moi trng nc, v du
nh mau.
Glycoprotein: co nhom them la cac momo- hoac oligosaccharide. Glycoprotein
co trong tat ca cac mo ong vat, thc vat va vi sinh vat. Thuoc nhom glycoprotein co
nhieu protien cua mau nh cac imunoprotein, fibrinogene, musin trong nc bot va
mang nhay, mot so enzyme (bromelain, ribonuclease B cua tuyen tuy), cac protein cau
truc cua mang te bao. Phan glucid cua glycoprotein cua mang te bao quay ra pha
ngoai cua mang te bao, va lam nhiem vu nh hng glycoprotein trong mang, va
ong vai tro nhan biet gia cac te bao. Cac goc glucid thng ket hp vi cac nhom
OH cua serine va threonine thong qua cac goc N-acetylglucosamine hoac N-
acetylgalactosamine.
Phosphoprotein: co nhom them la acid phosphoric, ket hp vi apoprotein qua
cac nhom OH cua serine va threonine cua protein. Phosphoprotein pho bien trong c
the sinh vat, tham gia ieu hoa nhieu qua trnh quan trong. Thuoc nhom nay co mot so
enzyme (nh phosphoglucomutase, phosphorylase A), casein cua sa, vitelin cua long
o trng ga.

Hoa sinh hoc - 31 -
Metaloprotein: Theo nh ngha chung cua protein phc tap, trong phan t
metaloprotein co cha kim loai, thong thng la Fe, Mg, Cu, Zn, Mo v.v
Metaloprotein co the co nhieu chc nang khac nhau nh van chuyen va d tr
kim loai, lien ket gia kim loai vi apopotein khong ben, trc tiep tham gia trong hoat
ong xuc tac cua enzyme.
Kieu phan loai protein th hai la chia protein thanh hai nhom hnh cau va hnh
si. Phan ln protein hnh cau tan c trong nc, c cau tao t mot hoac mot so
chuoi polypeptide. Cac ten goi albumin, globulin van con c dung nhng la e ch
nhng protein cu the, v du albumine trng, globulin huyet thanh v.v...
Protein si cung co the cha ng mot hoac mot so chuoi polypeptide. Chung la
nhng phan t co hnh dang keo dai, khong oi xng, chieu dai vt rat nhieu lan so
vi ban knh. Protein si chu yeu co mat trong cac mo lien ket, mo an hoi, mo co rut
vi chc nang cau truc, hoac la nhng chat khong tan cua toc va da.
VIII. PHAN GIAI PROTEIN.
Protein trong c the song chu yeu am nhan chc nang cau truc va xuc tac. Tuy
nhien, chung van c dung lam nguon nang lng. Trong trng hp nay trc tien
chung phai c thuy phan thanh aminoacid t do. Qua trnh thuy phan protein c
thc hien nh hai nhom enzyme peptidase. Endopeptidase co nhiem vu cat mach
polypeptide thanh nhieu oan oligopeptide phan t nho; trong khi o exopeptidase tac
ong len phan t protein t mot trong hai au tan cung cua chuoi polypeptide.
Thuoc nhom endopeptidase co pepsin, trypsin, chymotrypsin ong vat va
papain, bromelin cung nhieu enzyme khac thc vat.
Pepsin c tiet ra t mang nhay da day cua ong vat bac cao, co pH toi thch 1,0
1,5. Tien than cua no la dang khong hoat ong pepsinogen. No c hoat hoa thanh
pepsin nh HCl. Enzyme nay phan giai nhng lien ket peptide von hnh thanh t cac
aminoacid vong thm va dicarboxylic.
Trypsin va chymotrypsin do tuyen tuy tiet ra, thc hien tac dung xuc tac trong
ruot non vi pH 7-8. Tai o chung tiep tuc phan giai cac chuoi peptide cua thc an t da
day a xuong sau khi a c pepsin phan giai s bo. Nhng enzyme nay hnh thanh
t cac dang tien than khong hoat ong tng ng trypsinogen va chymotrypsinogen.
Trypsinogen c hoat hoa thanh trypsin nh enterokinase do thanh ruot tiet ra cung
nh nh chnh trypsin. Trypsin cung co tac dung hoat hoa chymotrypsinogen thanh
chymotrypsin. Trong tac dung xuc tac cua mnh trypsin phan giai cac lien ket peptide
co s tham gia cua nhom carboxyl cua arginine hay lysine. Trong khi o chymotrypsin
phan giai nhng lien ket peptide von hnh thanh vi s tham gia cua nhom carboxyl cua
tryptophan, phenylalanine hoac tyrosine.
Papain cua u u hoat ong trong pham vi kha rong t acid yeu en base yeu,
tuy thuoc vao ban chat cua c chat. No cong pha nhng lien ket peptide co leucine
glycine va cac diaminoacid tham gia. Cung nh cac loai peptidase thc vat khac, no
c hoat hoa nh HCN va cac hp chat cha nhom SH.

Hoa sinh hoc
- 32 -

Song song vi endopeptidase, cac chuoi polypeptide con b phan giai bi cac loai
exopeptidase. Carboxypeptidase phan giai chuoi polypeptide au C-tan cung,
aminopeptidase t au N-tan cung, lan lt tach ra cac phan t aminoacid t do.
Carboxypeptidase A cha kem (Zn
2+
) va thuy phan tat ca cac lien ket C-tan cung,
tr nhng lien ket ma aminoacid C-tan cung la lysine, arginine hoac proline.
Carboxypeptidase B ch tach t au C-tan cung cac aminoacid lysine va arginine.
Leucineaminopeptidase thuy phan cac lien ket peptide N-tan cung vi s tham
gia cua phan ln aminoacid.
Cac exopeptidase noi tren eu co mat trong ruot non. Cung vi endopeptidase
chung thuy phan protein thc an thanh aminoacid t do e c hap thu qua thanh ruot
vao mau.
C che cua qua trnh thuy phan protein noi bao cha c hieu biet nhieu, mac du
trong mot so mo va c quan, v du trong gan, no xay ra vi toc o rat nhanh.
IX. PHAN GIAI AMINOACID.
1.Chuyen amin hoa.
Nhom -amine cua aminoacid thng c tach khoi phan t giai oan au cua
qua trnh d hoa bang cac c che chuyen amin hoa hoac desamine hoa.
Nh phan ng chuyen amine hoa nhom -amine cua t nhat 11 aminoacid
(alanine, arginine, asparagine, cisteine, isoleucine, lysine, phenylalanine, tryptophan,
tyrosine, valine, va acid asparaginic) c tach khoi phan t va c mang en carbon
cua mot trong ba cetoacid la acid -cetoglutaric, acid pyruvic va acid asparaginic.
Ket qua la aminoacid bien thanh cetoacid, con cetoacid bien thanh aminoacid.
Phan ng chuyen amin hoa c thc hien nh cac enzyme aminotransferase
(hay transaminase). Trong so nhng enzyme nay c hieu biet nhieu nhat la alanie
trasaminase va glutamate transaminase xuc tac cac phan ng tng ng sau ay:
-Aminoacid + Pyruvate -Cetoacid + Alanine
-Aminoacid + acid -Cetoglutaric -Cetoacid + Acid glutamic
Transaminase co ca trong ty the va trong bao tng, nh o nhom -amine c
van chuyen gia hai c quan nay cua te bao. Coenzyme cua transaminase la
pyridoxalphosphate, dan xuat cua vitamine B
6
. Trong qua trnh phan ng xay ra s
chuyen hoa thuan nghch gia pyridoxalphosphate va pyridoxaminephosphate:

phan ng nay trai qua giai oan trung gian vi s hnh thanh cac hp chat
ketimine (hnh 11.1)
Hoa sinh hoc - 33 -

Hnh 11.1. Cac phan ng chuyen amin hoa

2. Desamin hoa.
Trong khi nhom -amine c loai khoi phan t aminoacid chu yeu bang con
ng chuyen amin hoa th cac nhom amine khac lai c loai khoi phan t chu yeu
bang c che desamine hoa. Trong mot so trng hp desamine hoa cung c s dung
cho nhom -amine. Desamine hoa la qua trnh loai bo nhom amine dang phan t
ammoniac. Kieu desamine hoa quan trong nhat la desamine hoa oxy hoa acid glutamic
vi s xuc tac cua glutamate dehydrogenase:
L-Glutamate + NAD
+
-Cetoglutarate + NH
4
+
+ NAD.H + H
+
Thong qua phan ng chuyen amine hoa nhom amine t cac aminoacid khac nhau
cuoi cung eu co the tr thanh nhom amine cua acid glutamic. V vay, phan ng tren la
con ng desamine hoa mot cach gian tiep moi aminoacid.
Hnh I.6. Cac phan ng chuyen amin hoa.
Glutamate dehydrogenase la mot enzyme ieu hoa, chiem v tr then chot trong
trao oi aminoacid. Hoat tnh cua no b c che bi ATP, GTP, NAD.H va c hoat hoa
bi ADP va nhieu aminoacid khac nhau. No cung chu anh hng cua tyroxine va mot
so hormone steroid.
mot so c the desamine hoa oxy hoa c thc hien nh hai dehydrogenase
khac ma nhom them la dan xuat cua flavine:
- Oxydase L-aminoacid co nhom them la FMN, xuc tac phan ng:
L-Aminoacide + FMN + H
2
O -Cetoacid + NH
3
+ FMN.H
2

- Oxydase D-aminoacid co nhom them la FAD, xuc tac phan ng tng ng oi
vi D-aminoacide. Tuy nhien, nhng enzyme nay ong vai tro khong quan trong lam

Hoa sinh hoc - 34 -
trong viec trao oi nhom amine. Ngi ta cha ro vai tro cua enzyme sau cung trong c
the ong vat, ni ma hau nh khong co D-aminoacid.
3. Decarboxyl hoa.
B decarboxyl hoa, aminoacid mat nhom COOH va bien thanh amine hoac
diamine. V du:
CH
3
CHNH
2
COOH CH
3
CH
2
NH
2
+ CO
2

H
2
N (CH
2
)
4
CHNH
2
COOH H
2
N (CH
2
)
5
CHNH
2
+
CO
2
(Cadaverine)
Amine la nhng chat rat oc oi vi c the ong vat cung nh thc vat. ong
vat chung hnh thanh trong qua trnh len men thoi, v du trong ruot gia. thc vat ngi
ta nhan thay cadaverine va putresine (san pham decarboxyl hoa cua ornitin) xuat hien
nhieu khi cay thieu kali. Trong trng hp o cay co the b chet.
Cac phan ng desamine hoa aminoacid c xuc tac bi nhom enzyme
decarboxylase ma nhom them cung la pyridoxalphosphate.
Trong c the cac amine chuyen hoa tiep tuc nh monoaminoxydase hoac
diaminoxydase:
O
R CH
2
NH
2
+ H
2
O +O
2
R- C + NH
3
+ H
2
O
2
H
thc vat amine co the tham gia tong hp cac hp chat d vong, trong o co
alcaloid
4. So phan cua ammoniac va chu trnh urea.
Mot lng ln ammoniac xuat hien trong qua trnh d hoa aminoacid va cac hp
chat nit khac. thc vat no c hap thu lai thong qua cac phan ng tong hp
aminoacid, muoi amon cua acid hu c hoac urea. ong vat co vu, lng the va mot
so loai ca urea cung c tong hp e sau o b bai tiet ra moi trng.
Viec tong hp amide (asparagine va glutamine) c thc hien nh asparagine
synthetase va glutamine synthetase nh nang lng cua ATP:
Acid asparaginic + NH
3
+ ATP Asparagine + ADP + Pvc
Acid glutamic + NH
3
+ ATP Glutamine + ADP + Pvc
Trong nhieu trng hp NH
3
tao muoi vi cac acid hu c trong dch bao cua
thc vat, v du vi acid malic, acid oxalic v.v...

Hoa sinh hoc
- 35 -

Tong hp urea thc vat va mot so ong vat la mot qua trnh gom nhieu giai
oan goi la chu trnh urea hay chu trnh ornitine (hnh 11.2).
Hnh 11.2 Chu trnh ornitine (chu trnh urea).

5. D hoa aminoacid va chu trnh acid tricarboxylic.
S phan giai aminoacid c thc hien bang nhieu con ng khac nhau, song
eu dan en mot so t san pham e sau o tiep tuc tham gia chu trnh acid tricarboxylic
(hnh 11.3).
Cac aminoacid khac nhau bang cac con ng khac nhau i vao chu trnh acid
tricarboxylic tai 5 iem: acetyl-CoA (qua pyruvate hoac qua acetoacetyl-CoA), -
cetoglutarate, succinyl-CoA, fumarate va oxaloacetate. Chi tiet cua nhng con ng
nay rat phc tap. Co the tm thay chung trong nhieu tai lieu giao khoa khac nhau ve hoa
sinh.

Hoa sinh hoc
- 36 -

Hnh 11.3. Moi lien he gia d hoa aminoacid va chu trnh acid tricarboxylic.
X. SINH TONG HP AMINOACID.
Cac nhom sinh vat khac nhau co kha nang tong hp aminoacid khong giong nhau.
Thc vat va nhieu loai vi sinh vat co the tong hp tat ca nhng aminoacid ma chung can
t ammoniac, nitrate hoac nitrite. Cay bo au va mot so thc vat khac, nh vi khuan not
san song cong sinh vi he re cua chung, co the s dung ca nit phan t e tong hp
aminoacid. ong vat thng ch tong hp c mot so aminoacid nhat nh. Nhng
aminoacid can thiet con lai (aminoacid khong thay the) can c tiep nhan t moi
trng qua thc an. Aminoacid khong thay the oi vi ngi la valine, leucine,
isoleucine, lysine, metionine, threonine, phenylalanine va tryptophan; trong nhng ieu
kien nhat nh ca histidine va arginine.
1.Kh nitrate va co nh nit.
Ammoniac la c chat trc tiep e tong hp aminoacid. Trong c the thc vat va vi
sinh vat no hnh thanh trong qua trnh kh nitrate hoac co nh nit phan t.
Qua trnh kh nitrate thanh ammoniac c thc hien qua hai giai oan chnh:
kh nitrate thanh nitrite nh nitrate reductase va kh nitrite thanh ammoniac nh nitrite
reductase. Nitrate reductase la mot flavoprotein cha molipden, s dung NADP.H lam
chat cho ien t. Trong qua trnh phan ng dong ien t c van chuyen theo trat t :
NADP.H FAD Mo NO
3
-

Hoat ong cua nitrite reductase can s tham gia cua feredoxin kh von hnh thanh
trong giai oan phan ng sang cua quang hp vi t cach chat cho ien t:
Feredoxin kh NAD FAD NO
2
-

Co nh nit phan t
c thc hien chu yeu
nh Rhizobium cong sinh
vi thc vat bac cao. Tao
lam, Azotobacter, vi
khuan quang hp va mot
so vi khuan song t do
khac cung co kha nang
nay. Qua trnh co nh
nit phan t c thc
hien nh hydrogenase,
feredoxin va hai loai
Hnh 11.4. S o dong ien t trong qua
h h i h
Hoa sinh hoc - 37 -
protein enzyme khac, mot
loai cha sat (Fe-protein),
loai kia cha sat va
molypden (Fe-Mo-
protein). Nang lng can
cho qua trnh nay do ATP
cung cap. S o dong ien
t trong qua trnh co nh
nit co dang khai quat nh
mo ta hnh 11.4.
Oxide carbon c che qua trnh nay bang cach hng dong ien t en H
+
e khoi
phuc hydro phan t cung nh ngan can viec van chuyen ien t t hydrogenase en
feredoxin.
He enzyme co nh nit khong ac hieu oi vi N
2
va co the kh ca oxide nit,
acetylene, cyanua va mot sochat tng t khac.
2. Amin hoa kh.
Acid glutamic c xem la aminoacid s cap ma t o tong hp nen hau nh tat
ca aminoacid con lai. No c hnh thanh trong phan ng amin hoa kh di tac dung
cua glutamate dehydrogenase:
NH
3
+ -Cetoglutarate + NADP.H + H
+
L-Gllutamate + NADP
+
.
Co the noi phan ng nay la nen tang oi vi qua trnh sinh tong hp aminoacid
trong moi c the. Nh phan ng chuyen amine hoa t acid glutamic se hnh thai cac
aminoacid khac. Rieng asparagine va glutamine c tong hp theo c che trnh bay
trong muc II.3.
Alanine va acid asparaginic c tong hp chu yeu bang c che chuyen amine
hoa, nhng chung cung co the hnh thanh trong qua trnh amine hoa kh. thc vat va
mot so vi sinh vat acid asparaginic con xuat hien nh phan ng amin hoa acid fumaric
do aspartate ammoniac lyase xuc tac:
HOOC CH = CH COOH + NH
3
HOOC CH
2
CHNH
2
COOH
Tuy nhien, trong te bao phan ng nay chu yeu hng ve pha giai phong
ammoniac.
3. Tong hp cac aminoacid th cap.
Tong hp cac aminoacid th cap la qua trnh rat phc tap, ac biet la oi vi cac
aminoacid khong thay the. Moi aminoacid thng c tong hp nh mot he thong a
enzyme hoat ong theo nguyen tac lien he ngc nham thc hien nguyen tac tiet kiem

Hoa sinh hoc - 38 -
von chi phoi moi hoat ong cua te bao. Nhieu tai lieu giao khoa ve hoa sinh hoc co the
giup chung ta tm hieu chi tiet van e nay.
XI. SINH TONG HP PROTEIN.
1. Cac yeu to can thiet cho sinh tong hp protein va cac giai oan cua
qua trnh nay.
Sinh tong hp protein, ma sinh hoc phan t goi la qua trnh dch ma (translation)
la mot qua trnh phc tap xay ra trong ribosome, ma nh ta a biet, c cau tao chu
yeu t protein va acid nucleic. Nguyen lieu e tong hp protein, tc aminoacid, c
tARN mang en ay, e di s ieu khien cua mARN va s xuc tac cua hang loat
enzyme tap hp thanh chuoi polypeptide vi thanh phan va trat t aminoacid a c
nh san trong cac gen tng ng. Nguon nang lng e tao ra cac ai phan t protein
la ATP hoac cac hp chat tng t san co mat trong te bao. Ngoai ra, trong cac te bao
nhan that viec tong hp hang loat protein con oi hoi s tham gia cua cac he thong cau
truc mang cua te bao ma trc het la he thong mang cua mang noi chat.
Toan bo qua trnh sinh tong hp protein co the c chia thanh 4 giai oan chu
yeu. o la: 1/ hoat hoa aminoacid; 2/ xay dng phc he m au; 3/ tang trng mach
polypeptide va 4/ ket thuc chuoi polypeptide.
Hoat hoa aminoacid la qua trnh bao gom hai bc;
a/ Aminoacid + ATP Aminoacyladenylate + PPvc;
b/ Aminoacyladenylate + tARN Aminoacyl-tARN + AMP.
Ca hai phan ng eu c xuc tac bi mot enzyme aminoacyl-tARN-synthetase
ac hieu cho moi aminoacid. Nh trong phan t enzyme cha 2 trung tam xuc tac, mot
ac hieu vi aminoacid va mot ac hieu vi tARN van chuyen aminoacid o, nen
enzyme cho phep aminoacid tm gan vi tARN ac hieu cua mnh.
Cong thc cau tao cua aminoacyladenylate va aminoacyl-tARN c trnh bay
trong hnh 11.5. Qua o ta co the thay nang lng cua ATP a c chuyen cho lien ket
ester giau nang lng gia nhom carboxyl cua aminoacid va nhom 3'-OH cua goc
adenylate tan cung cua tARN. Nh o aminoacyl-tARN tr thanh mot san pham hoat
ong, de dang tham gia phan ng polymer-hoa sau nay.

Hoa sinh hoc - 39 -

A B
Hnh 11.5. Aminoacyladenylate (A) va minoacyl-tARN (B)

e chuan b cho phan ng polymer hoa, cac phan t aminoacyl-tARN lan lt c a
vao khu vc A cua phan di n v ln cua ribosome, nh mo ta trong hnh 11.6. Tai
ay, nh ta se thay, no co ieu kien thuan li e lien ket vi chuoi polypeptide ang
tang trng von gan vi ribosome tai khu vc P.

Hnh 11.6. V tr cua aminoacyl-tARN trong ribosome

Xay dng phc he m au E. coli la mot qua trnh gom nhieu bc nh mo
ta trong hnh 11.7 vi s tham gia cua formylmethionyl-tARN
f
, va cac yeu to m au
IF
1
, IF
2
va IF
3
, von la nhng protein ac hieu.
Viec xay dng phc he m au trong cac te bao nhan that tren nhng net
chnh cung giong nh E. coli, song vi cac yeu to m au khac, ky hieu la eIF
1
, eIF
2

va eIF
3
. Them vao o, methionyl-tARN tham gia xay dng phc he m au khong b
formyl-hoa. Cac ribosome, nh ta a biet, cung co kch thc ln hn.


Hoa sinh hoc - 40 -

Hnh 11.7. Qua trnh hnh thanh phc he m au.
Tang trng chuoi polypeptide bao gom 3 bc chnh (hnh 11.8):
Bc 1: Sau khi fMet-tARN
f
gan vi ribosome tai khu vc P, aminoacyl-
tARN tiep theo ma codon cua no nam ke can codon cua fMet se i vao khu vc A cua
ribosome. Qua trnh nay can GTP e cung cap nang lng va mot loai protein co ten la
yeu to T, bao gom hai loai: Tu va Ts.
Bc 2: Nh peptidyltransferase, Lien ket peptide c hnh thanh gia nhom -
NH
2
cua aminoacid mi i vao vi nhom -COOH cua aminoacid ng trc. Dipeptide
mi xuat hien c gan vi tARN cua aminoacid th hai tai khu vc A, con tARN cua
aminoacid th nhat (bay gi c goi la tARNp) van nam lai khu vc P. Nang lng
can cho s hnh thanh lien ket peptide c nhan t lien ket ester cao nang cua
aminoacyl-tARN.
Bc 3: Peptidyl-tARN c chuyen t khu vc A sang khu vc P nh
translocase (yeu to G) va GTP. Yeu to G c a ra khoi ribosome. Ribosome di
chuyen mot oan e cho codon tiep theo tiep nhan khu vc A, tao ieu kien cho
aminoacid th ba tham gia phan ng polymer-hoa...
Qua trnh tang trng chuoi polypeptide se ket thuc khi mot trong cac codon
cham cau (UAA, UAG, UGA) i vao v tr oi dien vi khu vc A. Nh yeu to giai
phong R, peptidyl-tARN tach khoi ribosom; tARN tach khoi chuoi polypeptide;
ribosome 70S phan ly thanh cac phan di n v 30S va 50S e sau o lai tham gia
xay dng phc he m au cho sinh tong hp chuoi polypeptide khac.
Trong qua trnh sinh tong hp protein nhieu ribosome cung gan tren mot phan t
mARN, tao thanh phc he co ten la polyribosome hay polysome. Trong phc he nay
moi ribosome tong hp mot si polypeptide. Trong E. coli va cac te bao tien nhan khac

Hoa sinh hoc - 41 -
qua trnh dch ma co the bat au ngay khi phan t mARN con ang tang trng tren
khuon ADN.

Hnh 11.8. Cac bc tang trng chuoi polypeptide.
2. ieu hoa sinh tong hp protein; mo hnh operon va ly thuyet ieu
hoa cua Jacob va Monod.
Ly thuyet ieu hoa sinh tong hp protein c Jacob va Monod e xuat tren c
s nhng dan lieu thc nghiem ve hien tng cam ng va tran ap tong hp protein thu
c oi vi te bao E. coli. Nhng dan lieu nay cho thay rang sinh tong hp cac enzyme
-galactosidase (E
1
), -galactoside permease (E
2
) va thyogalactoside transacetylase
(E
3
) trong te bao E. coli cung c cam ng bi c chat cua chung la galactose. Tc la
nhng enzyme nay (enzyme cam ng) ch xuat hien trong te bao khi co mat lactose
trong moi trng dinh dng.
C che ieu hoa sinh tong hp cac enzyme cam ng c Jacob va Monod giai
thch nh sau thong qua v du cac enzyme d hoa lactose E
1
, E
2
va E
3
noi tren (hnh
11.9).
Ba enzyme c ma hoa bi cac gen cau truc X, Y va Z. Chung cung nhau tao
thanh mot operon (trong trng hp nay la lac-operon). Hoat ong cua operon nh mot
the thong nhat di s ieu khien cua mot gen khi ong O (operator). Ve phan mnh,
operator lai chu s ieu khien cua gen ieu hoa R (regulator) thong qua mot chat tran
ap co ban chat protein do gen nay tao ra. Khi gan vi operator, chat tran ap se lam cho

Hoa sinh hoc - 42 -
cac gen cau truc tren operon khong hoat ong, mARN khong c sao chep va do o
protein (tc 3 enzyme noi tren) khong c tong hp.

Hnh 11.9. ieu hoa sinh tong hp enzyme cam ng theo Monod va Jacov.
Khi chat cam ng allolactose (mot san pham cua lactose) gan vi chat tran ap,
lam cho no mat hoat tnh, tao ieu kien cho operon hoat ong. Tuy nhien, viec sao chep
thong tin t cac gen cau truc X, Y va Z ch thc s bat au khi cAMP cung vi mot loai
protein ac biet co ten la CAP tao thanh phc he hoat ong e gan vao khu vc
promoter nam ke can vi vi operator va goi mot phan len operator, giup chat cam ng
lam mat hoat tnh cua chat tran ap va ay no ra khoi khu vc nay.
S hnh thanh cAMP b c che bi mot san pham d hoa nao o cua glucose. V
vay khi co mat glucose trong moi trng dinh dng th cAMP khong c tao ra va
CAP t no khong the gan vao operator. V vay du co mat chat cam ng allolactose, cac
enzyme thuoc lac-operon van khong c tong hp.
Hoat ong cua operon tryptophan (Trp operon) ieu hoa tong hp cac enzyme
xuc tac cac qua trnh sinh tong hp tryptophan (hnh 11.10) cung tng t nh hoat
ong cua Lac operon, nhng co iem khac la bnh thng trong te bao thieu tryptophan
th chat tran ap trang thai khong hoat ong. Khi o operator m, s sao chep va
phien dch cac gene cau truc c tien hanh bnh thng, tong hp nen cac enzyme can
thiet e tao san pham cuoi cung la tryptophan. Nhng khi te bao a san xuat u
tryptophan, hoac khi them tryptophan vao moi trng nuoi te bao th tryptophan se ket

Hoa sinh hoc - 43 -
hp vi chat tran ap vi t cach la chat ong tran ap, bien no thanh dang hoat ong.
Chat tran ap hoat ong se lap tc ket hp vi operator, lam cho no b khoa lai va s sao
ma se b nh ch.

Hnh 11.10. Operon tryptophan (Trp operon)
Theo c che nay ham lng tryptophan trong te bao a kiem tra s tong hp nen
chnh no. o la moi lien he ngc trong s ieu hoa cac phan ng enzyme bang san
pham cuoi cung. ieu nay cho phep te bao thch nghi vi nhng thay oi ot ngot cua
moi trng cung nh tranh c nhng hao ton ve nguyen lieu va nang lng dung e
tong hp d tha mot chat nao o.
Sinh tong hp protein ong vat bac cao con co the c ieu hoa nh he thong
hormone. Mot so hormone co tac dung hoat hoa hoac c che enzyme adenylate cyclase
va do o the hien tac dung ieu hoa sinh tong hp protein thong qua tac dung chi phoi
ham lng cAMP trong te bao. Mot so hormone khac, ac biet la cac hormone steroid,
do co kha nang di chuyen xuyen qua mang, nen co the trc tiep can thiep vao c che
ieu hoa bang cach gan vi chat tran ap khong hoat ong e bien no thanh dang hoat
ong.


Hoa sinh hoc - 44 -


Hoa sinh hoc - 45 -
CHNG 2. EMZYME

Enzyme la nhng protein co chc nang chuyen hoa rat cao, lam nhiem vu xuc
tac hang nghn phan ng hoa hoc ma t o tao nen toan bo qua trnh trao oi chat
trong te bao va mo.
Ngay nay a xac nh c hang ngan enzyme khac nhau, trong o co hang
tram loai thu nhan c dang tinh the rat tinh khiet. Tuy vay, cac dan lieu di truyen
hoc cho thay rang con rat nhieu enzyme can phai c tiep tuc tm kiem.
Nhng phat minh mi ve s ieu khien cua qua trnh sinh tong hp enzyme
mc o gen, ve kha nang t ieu hoa cua cac he thong enzyme, ve vai tro cua enzyme
trong cac qua trnh sinh trng, phat trien va phan hoa v.v... la nhng thanh tu vo
cung to ln cua sinh hoc hien ai, lam nay sinh hang loat lnh vc nghien cu mi cua
enzyme hoc.
I. CAC BIEU THC DUNG TRONG ENZYME HOC.
e xac nh hoat tnh cua enzyme, ngi ta s dung cac bieu thc sau ay:
1. So chuyeu hoa = so mol c chat a chuyen hoa trong mot phut;
2. n v enzyme quoc te U = Lng enzyme xuc tac s chuyen hoa cua mot
mol c chat trong mot phut;
3. Hoat tnh ac hieu = Katal/ Kg protein hoat ong.
4. Hoat tnh phan t gam = Katal/Mol enzyme.
Katal la mot loai n v hoat tnh cua enzyme; no tng ng vi lng chat xuc
tac co the lam chuyen hoa 1 mol c chat thanh san pham trong 1 giay. Quan he gia
katal va U nh sau:
1 Katal (Kat) = 1 Mol/s = 60Mol/min = 60.10
6
Mol/s = 6 x 10
7
U;
1 U = 1 Mol/min = (1/60)Mol/s = (1/60) Kat = 16,67 nKat.
II. BAN CHAT HOA HOC CUA ENZYME.
Enzyme co the la protein n gian (enzyme mot thanh phan) hoac protein phc
tap (enzyme hai thanh phan).
Hoat tnh xuc tac cua nhieu enzyme ch phu thuoc vao ac iem cau truc cua
ban than protein-enzyme. Trong khi o oi vi mot so enzyme khac hoat tnh cua
chung phu thuoc vao s ton tai cua mot thanh phan nao o khong co ban chat protein
ma ngi ta goi la cofactor. Cofactor co the la ion kim loai hoac cac hp chat hu c
phc tap. Trng hp sau ac trng cho enzyme hai thanh phan, va khi o cofactor

Hoa sinh hoc - 46 -
c goi la coenzyme. oi khi hoat tnh xuc tac cua enzyme phu thuoc vao s ton tai
cua ca ion kim loai va coenzyme.
Cac cofactor noi chung eu ben vi nhiet o cao; trong khi o protein enzyme
neu b un nong se mat hoat tnh xuc tac. S lien ket gia protein enzyme va cofactor
co the co mc o ben vng khac nhau. Thong thng chung lien ket vi nhau bang
cac lien ket yeu va de dang tach khoi nhau bang phng phap tham tch. Tuy nhien
cung co nhng trng hp protein enzyme va cofactor gan vi nhau bang lien ket
cong hoa tr kha ben vng. Trong trng hp o cofactor c goc la nhom them, v
du trong cytochrome c hem vi t cach la cofactor gan kha ben vng vi protein
enzyme bang cac lien ket cong hoa tr thong qua cac goc cysteine, methionine va
histidine (hnh VI-1)
Cac phc he t nhien gia protein
enzyme va cofactor c goi la
holoenzyme. Thanh phan protein enzyme
khong hoat ong sau khi a b loai bo
cofactor c goi la apoenzyme.

Hnh VI.1. Cytochrome c
Trong phan t enzyme cac ion kim
loai co the thc hien mot trong hai chc
nang: hoac lam cau noi gia enzyme va
c chat, hoac trc tiep lam nhiem vu xuc
tac. Mot so enzyme cha kim loai c
gii thieu trong bang VI.1.
Cac coenzyme t ban than
chung thc hien chc nang xuc
tac thng vi t cach la chat
van chuyen trung gian ien t,
nguyen t hay nhom nguyen t
t mot hp chat nay sang mot
hp chat khac. Nh ket hp vi
apoenzyme hoat tnh xuc tac
cua chung tang len gap nhieu
lan va mang tnh ac hieu cao.
Bang VI.1.Mot so enzyme cha kim loai
Enzyme Kim loai
Alcoholdehydrogenase,
Carboanhydrase,
Carboxypeptidase

Zn
2+
Phosphohydrolase,
Phosphoptrasferase
Mg
2+
Arginase, Phosphoptrasferase Mn
2+
Cac loai cytochrome,
peroxydase,
catalase, Feredoxin
Fe
2+
hoac Fe
3+
Pyruvatephosphokinase K
+
Nhieu coenzyme la dan
xuat cua vitamine (bang VI.2).
Trong mot so trng hp khac
coenzyme co the la cac hp
chat quinon, hem, nucleotide
v.v...

Hoa sinh hoc - 47 -
Qua trnh bien hoa c chat xay ra tai trung tam hoat ong cua enzyme. Trung
tam nay thng co cau truc phu hp vi c chat thuoc cac phng dien ban chat hoa
hoc va cau hnh khong gian.
Trung tam hoat ong cua enzyme mot thanh phan do mot so goc aminoacid ac
biet cau tao nen; trong khi o nhng enzyme hai thanh phan trung tam hoat ong
c hnh thanh vi s tham gia cua coenzyme cung vi cac goc aminoacid nhat nh.
Bang VI.2. Coenzyme dan xuat cua vitamine
Coenzyme Vitamine Chc nang
Pyridoxal
phosphate
Piridoxine Chuyen amin hoa; decarboxyl hoa; rasemate
hoa
Thyamine-
pyrophosphate
Thyamine
(Vit. B
1
)
Decarboxyl hoa oxy hoa;
van chuyen nhom aldehyde
Coenzyme A Acid pantotenic Van chuyen acyl; phan giai va tong hp acid
beo trong ieu kien hieu kh
Acid
tetrahydrofolic
Acid folic Van chuyen cac nhom mot carbon
Biotin Biotin (Vit. H) Van chuyen CO
2
NAD
+
, NADP
+
Acid nicotinic Van chuyen H
+
, e
-
FMN, FAD Riboflavin Van chuyen H
+
, e
-
III. ONG HOC CUA CAC PHAN NG ENZYME.
Bat ky phan ng hoa hoc nao,v du phan ng A P, s d xay ra c la nh
mot phan nang lng trong so cac phan t A cha nang lng ln hn so phan t con
lai, lam cho chung ton tai trang thai hoat ong. trang thai nay de dang pha v mot
lien ket hoa hoc hoac tao ra mot lien ket mi e lam xuat hien san pham P. Nang
lng can e chuyen toan bo so phan t cua mot mol vat chat ieu kien nhat nh
sang trang thai kch ong c goi la nang lng hoat hoa. Nang lng nay can thiet
e chuyen cac phan t tham gia phan ng sang mot trang thai trung gian giau nang
lng tng ng vi nh cua hang rao hoat hoa (hnh VI.2).Toc o cua phan ng t le
vi nong o cua phan t trang thai trung gian nay.
Khi tang nhiet o nang lng chuyen ong nhiet cua phan t tang len, lam cho
so phan t co kha na ng at trang thai trung gian tang len. V the khi tang nhiet o len
10
o
, toc o cua phu hoa hoc tang len khoang hai lan (Q
10
= 2).
Khac vi tac dung cua nhiet o, chat xuc tac lam tang toc o cua phan ng
bang cach lam giam nang lng hoat hoa. S ket hp gia chat phan ng va chat xuc
tac lam xuat hien trang thai trung gian mi vi mc nang lng hoat hoa thap hn. Khi
san pham hnh thanh, chat xuc tac lai c giai phong trang thai t do.

Hoa sinh hoc - 48 -
Cac phan ng
enzyme cung tuan
theo nhng nguyen
tac chung cua ong
hoc cac phan ng
hoa hoc. Tuy nhien,
chung con co nhng
ac iem rieng. Mot
trong nhng ac
iem o la hien
tng bao hoa c
chat. nong o c
chat thap toc o cua
phan ng enzyme t
le thuan vi nong o
c chat. Nhng neu
tiep tuc tang nong
o c chat th toc o
phan ng tang cham
dan,

Hnh VI.2. Bien thien nang lng t do
trong cac phan ng hoa hoc.
va khi nong o c chat at mot gia tr nao o, toc o cua phan ng khong tang na.
Trong nhng ieu kien o nong o enzyme la yeu to quyet nh toc o phan ng.
Mac du hien tng bao hoa c chat ac trng cho moi enzyme, nhng gia tr cu
the cua nong o c chat la gia tr ac trng. Thong qua nghien cu van e nay
Michaelis va Menten nam 1913 a e xuat mot phng trnh dien ta toc o cac phan
ng enzyme va neu len mot so ly thuyet chung ve ong hoc cua qua trnh nay. Thuyet
nay ve sau a c Briggs va Haldans phat trien them.
Cac tac gia tren nhan thay rang trong cac phan ng enzyme trc tien enzyme E
tao ra phc he ES vi c chat S. Sau o ES se c phan giai thanh san pham P va
enzyme E t do.
Theo nh luat khoi lng, qua trnh o co the c mo ta nh sau:
k k
1
E + S ES E + P
3
k
2
k
4
trong o k
1
la hang so toc o phan ng hnh thanh ES t E va S; k
2
la hang so toc o
phan ng phan giai ES thanh E va S; k
3
la hang so toc o pha n ng phan giai ES thanh
E va P; k
4
la hang so toc o phan ng hnh thanh ES t E va P.
trang thai can bang toc o hnh thanh ES bang toc o phan giai phc he nay:
k
1
[E][S] k
2
[ES] = k
3
[ES] k
4
[E][P].

Hoa sinh hoc - 49 -
Bien oi phng trnh nay, ta co:
[ES](k
2
+k
3
) = [E](k
4
[P] + k
1
[S])
[ES] k
4
[P] + k
1
[S] k
4
[P] k
1
[S]
[E] k
2
+ k
3
k
2
+k
3
k
2
+k
3
Do cac giai oan au cua phan ng gia tr cua [P] vo cung nho nen co the gian
lc phng trnh tren nh sau:
[ES] k
1
[S]
[E] k
2
+ k
3

at [E]
t
la ham lng enzyme tong so va K
m
= k
2
+k
3
/ k
1
, ta co:
[E] [E]
t
-[ES] [E]
t
K
m
[ES] [ES] [ES] [S]
Toc o ban au v cua phan ng enzyme t le thuan vi ham lng enzyme hoat
ong, hay [ES], nen ta co the viet:
v = k
3
[ES]
Neu nong o c chat rat ln, lam cho hau het enzyme trong he thong eu ton tai
trang thai ES, th toc o phan ng enzyme se at gia tr toi a V, va toc o toi a o
se bang:
V = k
3
[E]
t
Do o:
[E]
t
V K
m
[ES] v [S]
Nhan hai ve cho [S] va bien oi phng trnh, ta co:
V[S]
v =
K
m
+ [S]
ay chnh la phng trnh Michaelis-Menten va K
m
c goi la hang so
Michaelis.
Y ngha thc tien cua hang so Michaelis la cho no chnh la gia tr cua nong o
c chat khi toc o phan ng bang toc o toi a. Thay V va v bang cac con so tng
ng 1 va 0,5 vao phng trnh tren, ta se thay ro ieu o. Nh vay, K
m
c o bang
n v nong o, tc mol/l.
Hang so Michaelis la mot hang so rat quan trong. No xac nh ai lc cua enzyme
vi c chat. K
m
cang nho th ai lc nay cang ln, toc o phan ng cang cao v toc o
toi a V at gia tr nong o c chat cang thap.
Tren c s phng trnh Michaelis-Menten, bang cach xay dng ng bieu
dien s phu thuoc cua v vao [S] va bang o th o xac nh toc o toi a V ta co the
tm thay gia tr cua [S], o v = V/2, tc gia tr cua K
m
(hnh VI.3).

Hoa sinh hoc - 50 -
Tuy nhien, bang cach nay kho xac nh v mot cach chnh xac. e khac phuc
nhc iem o, ngi ta s dung ng bieu dien Linewear-Burk. Hai tac gia nay
bien oi phng trnh Michaelis-Menten thanh dang:
1/v = K
m
/V x 1/[S] +
1/V

Hnh VI.3. ng bieu dien
phng trnh Michaelis-Menten
u iem cua
phng 1trnh nay
la cho gia cac
ai lng 1/v va
1/[S] co moi lien
he t le thuan (hnh
VI.4).
Qua ng
bieu dien nay ta co
the thay rang tang
ABO = K
m
/V va
BO = 1/K
m
.
Phng trnh
nay con cho phep
tm hieu nhieu kha
canh quan trong
lien quan en tac
dung cua cac chat
c che hoat tnh
cua enzyme.
1/v

A
1/V
B
-1/Km O -1/[S]
Hnh VI.4. ng bieu dien
phng trnh Lineweaver-Burk
IV. ANH HNG CUA pH LEN HOAT TNH ENZYME.
Phan ln enzyme c ac trng bi s phu thuoc cua chung vao pH, trong o
moi enzyme co hoat tnh cao nhat mot gia tr pH nhat nh. ca hai pha cua gia tr
nay hoat tnh enzyme eu giam. Gia tr pH toi thch nay (pH
opt
) bien thien trong pham
vi kha rong, t 1,5-2,5 oi vi pepsin en 8-9 oi vi trypsin. V vay trong moi nghien
cu enzyme pH can c gi vng bang dung dch em thch hp.
S phu thuoc cua hoat tnh enzyme vao pH c qui nh bi pK cua cac nhom
ion hoa trong phan t enzyme, ac biet cua cac nhom tai hoac gan trung tam hoat
ong (co the co vai tro trong viec ket hp gia apoenzyme va coenzyme), hoac cac
nhom co the ong gop vao viec lam bien oi trung tam hoat ong thong qua viec lam
bien dang cac bo phan cua phan t enzyme. pH cung co the anh hng xa hn en
mc o ion hoa hoac cau truc khong gian cua c chat.

Hoa sinh hoc
- 51 -

Tuy nhien, co nhng trng hp ma hoat ong cua enzyme khong phu thuoc vao
pH cua moi trng. o la trng hp enzyme tac dung len nhng c chat trung hoa ve
ien hoac nhng c chat ma cac nhom tch ien khong co vai tro quan trong oi vi
tac dung xuc tac. Papain la mot trng hp nh vay.
Gia tr pH toi thch oi vi hoat tnh xuc tac cua enzyme khong nhat thiet phu
hp vi gia tr pH cua dch bao ma trong o cha enzyme. ieu o cho phep chung ta
ngh rang anh hng cua pH len hoat tnh enzyme la mot trong thng yeu to gop phan
ieu hoa hoat tnh enzyme trong te bao.
V. ANH HNG CUA NHIET O LEN HOAT TNH ENZYME.
Nhiet o anh hng len toc o cua moi phan ng hoa hoc. Trong pham vi bien
o sinh ly toc o cua phan ng enzyme tang theo nhiet o vi he so Q
10
bang 2. Khi
nhiet o vt qua gii han nhat nh, do protein enzyme b bien tnh, toc o cua phan
ng enzyme giam mot cach ot ngot. Nhiet o toi thch (t
o
opt
) cua phan ng enzyme
thng nam trong pham vi 40-50
o
C va co the bien oi phu thuoc vao nhieu yeu to nh
o sach cua enzyme, thi gian phan ng v.v...
Moi enzyme chu anh hung cua nhiet o mot cach khac nhau. ac iem o
c li dung e tach chiet va nghien cu hoat tnh cua tng loai enzyme trong te bao.
VI. HOAT HOA ENZYME.
Nhieu enzyme e the hien hoat tnh cua mnh, can phai co s ho tr cua cac yeu
to khac nhau, trong o moi enzyme c hoat hoa bang mot con ng nhat nh. Bon
kieu hoat hoa khac nhau c mo ta trong hnh VI.5.
Hoa sinh hoc - 52 -
Hnh VI.5. Cac kieu hoat hoa enzyme
Mot so enzyme, v u hoa bang cach cat
bo mo
nh thanh cau disulfide, v du
ribon
d pepsinogen, trypsinogen ... c hoat
t oan oligopeptide khoi phan t proenzyme (1);
Mot so enzyme khac c hoat hoa bang cach h
uclease (2), hoac bang cach tao pha vi ion kim loai (3). Kieu hoat hoa th t
ac trng cho cac enzyme d lap the, c thc hien bang cach thay oi cau hnh
khong gian cua enzyme nh mot effector dng tnh ac hieu (4).
VII. C CHE ENZYME.
Hoat tnh cua enzyme b c che khi protein enzyme b bien tnh di tac dung
cua k
hoat tnh cua enzyme nay nhng lai kch
thch
ac hieu oi
vi m
nghc
chat c che ong vai tro quan trong trong lnh vc y hoc. Nhieu cong trnh
nghie
vai tro rat quan trong trong viec nghien cu cac c
che ti
ngi ta chia chung lam hai nhom:
chat
ac hieu tuyet oi cua nhieu
phan
ien hnh la acid malonic. o la mot
chat c che manh va ac hieu oi vi enzyme suxinate dehydrogenase. Enzyme nay
im loai nang va cua nhng tac nhan pha v lien ket hydro, khi trung tam hoat
ong cua enzyme b phong toa hoac b bien dang di tac dung cua cac yeu to lat ly
hoac cac chat hu c va vo c khac nhau.
Mot hp chat hoa hoc co the c che
hoat tnh cua enzyme khac. V du, CN
-
c che nhieu enzyme oxy hoa kh
nhng lai kch thch hoat tnh cua mot so loai enzyme thuy phan (papain, catepsin...).
Mat khac, mot enzyme co the b c che bi mot hp chat hoa hoc mot nong o nao
o nhng lai chu tac dung kch thch cua hp chat ay mot nong o khac.
Ben canh nhng chat c che chung, co nhng chat ch gay c che
ot enzyme hoac mot nhom enzyme xac nh.
Tac dung c che oi vi hoat tnh enzyme co the thuan nghch hay khong thuan
h.
Cac
n cu c thc hien nham s dung chung e cha benh. Cac nha khoa hoc ac
biet lu y en van e s dung phng phap c che chon loc cac phan ng enzyme
nham ieu tr cac benh ung th.
Cac chat c che cung ong
nh vi cua trao oi chat. V du, cac chu trnh Krebs, Calvin...a c tm ra tren
c s s dung cac chat c che ac hieu khac nhau.
Tren c s c che tac dung cua chat c che
c che canh tranh va chat c che khong canh tranh.
c che canh tranh la hau qua cua s vang mat tnh
ng enzyme. Trung tam hoat ong cua enzyme co the ket hp vi cac chat c
che ma cau tao hoa hoc cua chung tng oi giong vi c chat, do o can tr s tiep
xuc cua c chat vi enzyme tai trung tam nay.
Mot trong nhng chat c che canh tranh

Hoa sinh hoc - 53 -
khong
h vi enzyme. Chat c che I ket hp vi enzyme E, tao thanh phc he EI, He so
phan
Trong phan ong hoc cac phan ng enzyme ta a thay rang:
[ES] [S]
V [E] = [E] [EI] [ES],
t
[EI] [ES] K
m
=
[ES] [ES] [ES] [ES] [S]
Hay:
[E]
t
K
m
[EI]

m
[E] [I]
Thay [EI] ba
K [ES]
t

[S]
V toc o toi a c o tc thi v t le
thuan vi nong o ph

.

phan biet c acid malonic (HOOC-CH
2
-COOH) vi acid suxinic (HOOC-
CH
2
- CH
2
-COOH). Khi trung tam phan ng cua suxinate dehydrogenase b acid
malonic chiem cho, enzyme nay khong con phan ng oxy hoa acid suxinic. Mc o c
che trong trng hp nay phu thuoc vao t le malonate/suxinate. Nong o cua acid
suxinic cao en mc nat nh se co the hoan toan loai tr tac dung c che cua acid
malonic. ac iem cua c che canh tranh la toc o cua phan ng phu thuoc vao nong
o chat c che, nong o c chat va ai lc cua enzyme vi c chat cung nh vi chat c
che.
Cac chat c che canh tranh thuoc nhom nhng chat c che co phan ng thuan
nghc
li K
i
cua phc he nay c goi la hang so c che.
[E][I]
K
i
=
[EI]
[E] K
m
=
ta co:
t
[E]
t
[EI] [ES] [E]
= - -
= + + 1
[ES] [S] [ES]
[E] [I] [E]
[ES] [S] K
t
K
ng , ta co: = + + 1.
i i
[E] [S] 1 K
m
V [ES] = , ta co the thay bang , tc:
K [ES] [E] [S]
m

[E]
t
K
m
K
m
[I]
= + + 1
[ES] [S] K
i
t
, con to V thuan vi enzyme tong so [E] t le
c he [ES], nen:
[E]
t
V K
i
K
m
K
m
[I] Ki[S] K
i
K
m
+ K
m
[I] + Ki[S]
= = + + =
[ES] v K
i
[S] K
i
[S] Ki[S] Ki[S]

Hoa sinh hoc - 54 -
V[S]

han r phng trnh Michaelis-
Menten cho K
m
c nhan vi he so (1 + [I]/K
i
).
qua va toc o phan ng se at gia
tr toi
nang cua chat c che tac dung vi enzyme, tc xac nh hang so K
i
. e giai
quyet
. +
[S] V
trng hp
khong co chat c che, ta se thay chung ch khac nhau gia tr (1+[I]/K
i
g cua
goc n
tranh la mot qua trnh c
che t
V[S]Ki
T o, v = =
K
m
K
i
+ K
m
[I] + K
i
[S] K
m
+ K
m
[I]/Ki + [S]

V[S]
v Tc: =
K
m
(1 + [I]/K
i
) + [S]

Ta e ang phng trnh tren khac vi co th n thay
V vay, c che canh tranh lam tang gia tr cua K
m
. Neu [S] co gia tr rat ln so
vi gia tr cua K
m
(1+[I]/K
i
) th gia tr nay co the bo
a. Neu K
i
nho va [I] ln th se xay ra s c che va mc o c che phu thuoc
vao [S].
Y ngha thc tien cua viec nghien cu tac dung c che la anh gia ve mat nh
lng kha
cong viec nay se thuan tien hn neu ap dung phng trnh Lineweaver-Burk
trong trng hp c che canh tranh vi dang sau ay:
1 [I] K
m
1 1
= (1 + )
v K

i
V

So sanh phng trnh nay vi phng trnh Lineweaver-Burk trong
), tc tan
ghieng bang (1+[I]/K
i
).K
m
/V, con gia tr 1/V hay V khong thay oi. ng bieu
dien cua hai phng trnh Lineweaver-Burk trong cac trong hp khong co va co c
che canh tranh c mo ta trong hnh VI.6.
c che khong canh

Hnh VI.6. ng bieu dien phng trnh
Lineweaver-Burk trong cac trng hp khong co
va co c che canh tranh.
huan nghch ma trong
o chat c che khong ket
hp vi enzyme tai v tr
von xay ra phan ng gia
enzyme va c chat. Trong
trng hp nay khong the
loai tr tac dung c che
bang cach nang cao nong
o c chat, v c chat khong
ngan can s ket hp cua
chat c che vi enzyme.
Mc o c che phu thuoc
vao [I] va K
i
.

Hoa sinh hoc - 55 -
Cac chat c che khong canh tranh thng co cau tr
vi c chat.
E + I EI
E]
t
[EI])

i
[EI]
Toc o v t ng rng le thuan vi [E]
t
, con toc o v
i

trong trng hp o c
i
[E]t [EI]
1

=
=

[E]

K
i
+
i

K
i
+
i
+ [I]
phng trnh tren, ta thay ro toc o phan ng khi co mat chat c che khong
canh tranh ch phu thu phu thuoc vao [S]. Toc o toi a V se b
giam
K
m
1
V[S] V
1 [I] K
m

+ , +
)
uc khong gian khong giong
Phng trnh cua c che khong canh tranh co the dien at nh sau:

[E] [I] ([
[I]
K = =
i
[EI] [EI]

K
i
[EI] = [E]
t
[I] [EI] [I]
K + [EI] [I] = [E]
t
[I]
[EI](K
i
[I]) = [E]
t
[I]
[E]
t
[I]
[EI] =
K
i
+ [I]
ro t hp khong co c che t
c hat c che t le thuan vi [E].
v [E]
t

Do [E] = [E]
t
[EI], nen = . Thay gia tr cua [EI], ta co:
v

v [E]
t
1
1
= =
vi

t

[I] [I] K [E]
t
[I] [I]
1
[I] K + [I] K + [I] K
i i

vi Ki
v Ki + [I]
T
oc [I] va K
i
ma khong
v mot phan enzyme khong tham gia phan ng c na. Neu K
i
nho, va [I] ln
th t le vi/v se rat nho. Trong trng hp ngc lai, t so o se gan bang 1, co ngha la
tac dung c che khong ang ke.Chuyen phng trnh tren sang dang phng trnh
Lineweaver-Burk:
1 K
i

v v
i
(Ki +[I])

Thay 1/v bang + , ta co:

K
i
K
m
1
1
= hay = (1 + ) (

Hoa sinh hoc - 56 -
v
i
(K
i
+ V v [I]) V[S]
V

Qua phng trnh tren, ta thay ro ca goc nghieng K
gia
tr bang (1 + [I]/K
i
).
phng trnh tren c
g cho
cac e
ng au mot he
e e
c che d lap the co the thuan nghch hay khong thuan
d lap the, no do mot effector am tnh gay ra. Chat nay hoat
khong canh tranh, tc lien ket vi enzyme tai mot trung tam khac vi trung tam hoat
ong,
co dang ch S (hnh
VI.8)
ng cua he thong
a en
i
K
i

V[S]
m
/V va 1/V eu tang len mot
ng bieu dien cua
mo ta hnh VI.7.
Kieu c che th ba
la c che d lap the. Kieu
c che nay ac trn
nzyme d lap
the hay
enzyme ieu hoa.
Nhng enzyme loai nay
thng
thong a nzym chu
trach nhiem xuc tac cho
mot loat phan ng gan
lien nhau ve mat chc
nang.
nghch. Trai vi hoat hoa
ong nh mot chat c che
lam cho cau truc khong gian cua enzyme bien oi va hnh dang cua trung tam
hoat ong khong con phu hp vi c chat na. Tuy nhien, trong trng hp nay s phu
thuoc cua toc o phan ng vao nong o c chat va chat c che khong n gian va
khong the mo ta bang s bien oi cua K
m
.
ng bieu dien
s phu thuoc nay thng

Hnh VI.7. ng bieu dien phng trnh
Lineweaver-Burk trong cac trng hp khong co va
co c che canh tranh.

Hnh VI.8. ng bieu dien s phu thuoc cua
toc o phan ng enzyme ieu hoa (d lap the)
vao nong o c chat.
.
Chat c che d lap
the thng la san pham
cuoi cu
zyme (nguyen tac
lien he ngc). Trong
phan trao oi chat ta se
e cap en nhieu trng

Hoa sinh hoc - 57 -
hp cu the cua kieu c
che nay.
VIII. TNH AC HIEU CUA ENZYME.
Moi enzyme ch tac dung len mot c chat nhat nh hay mot so c chat co cau
et hoa hoc nhat nh trong phan t
oi vi c chat.
c, tham ch rat giong vi c chat nay, chang han nh ester
meth
loai triacylglycerine khac nhau; phosphatase co the phan giai
nhieu
canh yeu to th nhat, c chat con cha mot hoac mot so nhom chc co
kha nang ket hp vi
tam h
ien hnh la trng hp
tao gan nhau, hoac ch tac ong len mot kieu lien k
c chat.
Nhng thay oi rat nho trong cau truc hoa hoc cua c chat oi khi co the lam
mat kha nang xuc tac cua enzyme. ac iem nay c goi la tnh ac hieu cua
enzyme
Nhng enzyme co tnh ac hieu hep, hay tnh ac hieu tuyet oi ch tac ong len
mot c chat nhat nhat nh. V du, arginase ch phan giai arginine ma khong phan giai
bat ky hp chat nao kha
ylic cua arginine.
Nhng enzyme co tnh ac hieu rong hay tnh ac hieu tng oi co the tac
dung len cac c chat khac nhau nhng co cung mot kieu lien ket hoa hoc.. V du lipase
co the phan giai nhieu
loai ester cua acid phosphoric.
Nhng enzyme co tnh ac hieu lap the ch co kha nang phan giai mot trong hai
dang ong phan quang hoc, v du dang D- hoac dang L-.
Co hai c s cau truc quan trong xac nh tnh ac hieu cua enzyme oi vi c
chat. o la:
1/ C chat can cha kieu lien ket hoa hoc ac trng ma enzyme co the cong pha;
2/ Ben
enzyme bang cach nao o e nh hng c chat tai trung
oat
ong, tc trung tam phan
ng, cua enzyme. V du
acetylcholine esterase
phan giai lien ket ester
gia choline va goc acetyl
(hnh VI.9). Kha nang cua
enzyme phan giai lien ket
ester phu thuoc ca vao s
ton tai cua cac goc serine,
tyrosine va histidine von
trc tiep tham gia qua

Hnh VI.9. S phu hp ve cau truc
gia enzyme va c chat
Hoa sinh hoc - 58 -
trnh phan ng, cung nh
vao s co mat cua nhom
COO
-
e lien ket tnh ien
vi N
+
cua c chat.
IX. DANH PHAP VA PHAN LOAI ENZYME.
Ten goi cua enzyme thng la ten goi cua c chat hay cua kieu phan ng ma no
. Ngoai ra con co nhng ten
t hoa hoc cua phan ng do
enzym
nay toan bo enzyme c chia thanh 6 nhom ln; moi nhom ln nay lai c
chia
c ky hieu
bang
at nhan ien t la NAD
hoac
4.1.1.1 Pyruvate
de
hetase
Ba
xuc tac cong vi uoi ase, v du urease,, hydrolase v.v..
goi truyen thong theo thoi quen, khong cho thay ban cha
e xuc tac, v du pepsin, trypsin ... ca hai kieu goi ten neu tren eu thieu chnh
xac.
e khac phuc tnh trang o, Hoi Hoa sinh hoc quoc te e ngh s dung mot he
thong danh phap va phan loai tren c s ban chat cua phan ng c xuc tac. Theo he
thong
thanh nhieu phan nhom; moi phan nhom nay lai c chia thanh nhieu phan
nhom nho hn, trong o bao gom nhng enzyme co c chat tac dung giong nhau. Moi
nhom, moi phan nhom va moi enzyme c ky hieu bang mot ma so a trng gom
tng ng mot, hai, ba hoac bon con so cach nhau bang cac dau cham.
Ten goi cua 6 nhom enzyme va cac phan nhom quan trong c gii thieu trong
bang II.3. cung vi ban chat cua cac phan ng c xuc tac.
Cac phan nhom nho hn thuoc moi phan nhom trong bang VI.3
nhng ma so gom 3 con so. V du ba phan nhom nho au tien cua phan nhom 1.1
la 1.1.1, 1.1.2 va 1.1.3 ac trng cho cac trng hp ma ch
NADP, FAD va cytochrome.
Ma so cua moi enzyme gom 4 con so, v du:
1.1.1.29 Glycerophosphate dehydrogenase; 2.7.1.1 Hexokinase
3.2.1.20 - - Glucosidase;
carboxylase;
5.3.1.1 Triosophosphate isomerase; 6.3.1.2 Glutamin synt
ng VI.3. Danh muc ma so cua 6 nhom enzyme va cac phan nhom chnh cua chung
Nhom Phan nhom Phan ng c xuc tac
1. Oxydoreduc

1.4

=C=O
DPH
tase Hydrogen hoa va dehydrogen hoa
1.1 =CHOH

1.2
1.3
1.5
1.6
CH=C
CHNH
H
2
=CHNH
, NA NADH

2. Transferase cac nhom chc
oac cetone

2.1
2.2
Van chuyen
Cac goc 1 carbon
hom aldehyde h N

Hoa sinh hoc - 59 -
2.3
2.4
hoac aryl
nh
2.5
2.6
2.7
2.8
Acyl
Lien ket glycoside
Nhom methylalkyl
Nhom cha nit
cha phosphore Nhom
Nhom cha lu huy

3. Hydrolase

3.2
3.3
3.4
n
lycoside
n ket C-N khac
it acid

3.1
3.5
3.6
Cac phan ng thuy pha
Ester
G
Eter
Peptide
Cac lie
Cac anhydr
4. Liase ket oi
4.1
4.2
4.3
Tao lien
=C=C=
=C=O
=C=N
Bang II.3. Danh muc ma so cua 6 nhom enzyme ung
iep theo)
5.2
5.3
5.4
Xis-trans-isomerase
t
han t
va cac phan nhom chnh cua ch
(t
5. Isomerase
5.1
an hoa
Rasemase va epimerase
ong ph

Oxy hoa noi phan
Transferase noi p
6. Ligase
6.4
6.1
6.2
6.3
Tao ra lien ket nh ATP
C=O
CS
=C=N
CC
X. HE THONG MULTIENZYM VA VAI TRO CUA ENZYME IEU
HOA.
c la c chat cua phan ng sau. Tuy thuoc vao mc o phc tap, nhng he
thong multienzyme (hay a enzyme) nay
ong oc
lap nhau. Cac phan
kch
hp nhau thanh mot phc he
hoat ong phoi hp nhau. o la
Trong te bao nhieu enzyme hoat ong ong thi, trong o san pham cua phan
ng tr
c chia lam 3 loai (hnh VI.10):
1/ Cac enzyme ca biet hoa tan
trong te bao chat va hoat
t c chat co
thc nho, de khuech tan, co
the tm thay nhanh chong con ng
t enzyme nay sang enzyme khac
(1);
2/ cac enzyme cua he thong
ket

Hnh VI.10. Cac kieu to chc cua
he thong multienzyme
Hoa sinh hoc - 60 -
trn
3/ Co mc o to chc cao nhat la nhng he
truc tren phan t cua te bao. o la trng hp
chuyen ien t trong ti the. Moi enzyme ca biet gan vao lp mang trong cua ti the,
va la
n mot
cach
Trong a
so trng hp
san pham cuoi
cung co tac
ne

bao
g hp he enzyme tong hp acid
beo cua nam men. He thong nay
gom 7 enzyme ket hp chat che vi
nhau. Neu tach ri nhau, tat ca eu
b mat hoat tnh;
thong enzyme lien ket vi cac cau
oi vi he enzyme cua chuoi van
m nhiem vu xuc tac, va ong vai tro nh mot yeu to cau truc cua mang.
Trong nhng he thong multienzyme nay toc o cua mot phan ng nao o thng
xac nh toc o cua toan bo he thong, trong o yeu to han che co the la nong o
enzyme hoac nong o c chat. Viec ieu hoa toc o nay co the c thc hie
t ong.

Hnh VI.11. c che L-treonine
desaminase bi isoleucine theo
nguyen tac lien he ngc.
dung c che
enzyme au
tien cua he
thong, nh o
toc o cua
toan bo qua
trnh cung
c xac nh
bi trang thai
nong o cua
san pham cuoi
cung. V du he
thong
multienzyme,
trong o L-
threoni
chuyen hoa
thanh L-
isoleucine
gom 5 phan

Hoa sinh hoc - 61 -
ng (hnh
VI.11).
cuoi cung L- Enzyme E
1
(threonine desaminase) b c che bi san pham
isoleucine, mac du no khong phai la chat canh tranh vi L-threonine. Tat ca nhng
amino
oa.
tnh, trong o moi effector
ket h
oi polypeptide, trong phan t cua chung ben canh mot hoac
mot s
tac dung lam bien oi cau hnh khong
gian c
acid giong no eu khong gay hieu ng nay. Ro rang la khi trong he thong tch
luy qua nhieu L-isoleucine, vt qua mc cho phep nao o, th enzyme au tien cua
he thong se b c che. Kieu c che nay c goi la c che theo nguyen tac lien he
ngc. Enzyme b c che bi san pham cuoi cung trong he thong multienzyme c
goi la enzyme ieu hoa (hay enzyme di lap the). Chat trao oi gay tac dung c che
enzyme ieu hoa c goi la effector am tnh, hay modulator am tnh. Cung co trng
hp enzyme ieu hoa nam iem phan nhanh cua cac day phan ng. Enzyme ieu
hoa co the la n tr, neu ch chu tac dung cua mot effector, hoac a tr neu chu tac
dung cua t hai effector tr len. Cac effector co the la cac san pham cuoi cung cua
nhng trat t phan ng lien quan nhau. Nh ac iem nay mot so he thong
multienzyme co the co chung mot he thong ieu hoa.
Enzyme ieu hoa co the chu tac dung cua effector dng tnh. Thong thng,
effector dng tnh la c chat cua chnh enzyme ieu h
Co nhng trng hp enzyme ieu hoa chu tac ong ong thi cua mot hoac
mot so effector am tnh va mot hoac mot so effector dng
p vi be mat enzyme tai mot v tr rat ac trng.
Phan ln enzyme ieu hoa trong ieu kien te bao la nhng enzyme xuc tac cac
phan ng khong thuan nghch.
S ton tai cua enzyme ieu hoa la mot trong nhng the hien cua nguyen tac tiet
kiem toi a cua te bao.
Enzyme ieu hoa thng co kch thc rat ln va cau truc rat phc tap, c
hnh thanh t nhieu chu
o trung tam hoat ong, hay trung tam c chat dung e thc hien nhiem vu xuc
tac con co nhng trung tam dung e ket hp vi effector am hoac dng tnh goi la
trung tam ieu hoa hay trung tam d lap the.
Tac dung ieu hoa bang cach gay hieu ng am cung nh dng, theo quan iem
pho bien hien nay, c thc hien thong qua
ua enzyme, lam cho hoat tnh xuc tac cua enzyme tang hoac giam.
XI. ISOENZYME
Nhieu enzyme tr
nhieu dang phan t kha
ong cung mot c the, tham ch trong cung mot te bao, ton tai
c nhau. Nhng dang phan t o cua cung mot enzyme c goi
la isoenzyme. Chung thng c tach khoi nhau mot cach de dang bang phng phap
ien di.

Hoa sinh hoc - 62 -
V du lactate dehydrogenase trong cac mo cua chuot bach co 5 dang isoenzyme
ma ngay nay a tach c dang tinh khiet. Chung eu xuc tac cho mot phan ng nh
nhau,
H. Mot isoenzyme chiem u the trong c c cau tao t 4 chuoi M
giong
g moi loai isoenzyme c kiem tra mc o gen va co
the bi
quan. Ngi ta cho rang khong nhng
enzym
song gia tr K
m
cua chung rat khac nhau. Ca 5 dang nay eu co trong lng phan
t vao khoang 134.000 va cha 4 chuoi polypeptide (trong lng phan t cua mot
chuoi la 33.500).
5 isoenzyme la 5 kieu phoi hp khac nhau cua hai loai chuoi polypeptide
chuoi M va chuoi
het nhau (ky hieu la M
4
). Isoenzyme th hai chiem u the trong tim, ky hieu la
H
4
, c hnh thanh t 4 chuoi H. Ba isoemzyme con lai, ky hieu la M
3
H, M
2
H
2
va
MH
3
. Ngi ta a tach rieng c cac chuoi M va H. trang thai nay chung khong
con kha nang xuc tac. Hai loai chuoi co thanh phan va trat t aminoacid khac nhau.
Khi tron trong ong nghiem hai loai chuoi vi t le tng ng cac kieu isoenzyme khac
nhau se t ong xuat hien.
S tong hp cac chuoi M va H c ma hoa bi hai gen khac nhau, Nh vay, t
le tng oi cua chung tron
en oi trong qua trnh phat trien cua phoi.
Viec nghien cu isoenzyme co y ngha rat quan trong oi vi viec tm hieu c s
phan t cua s phan hoa te bao va phat sinh c
e ma nhieu loai protein cung ton tai trong te bao cac dang khac nhau, phan
biet nhau bi t le gia cac chuoi polypeptide von c ma hoa bi cac gen khac nhau.


Hoa sinh hoc - 63 -
CHNG 3. KHAI NIEM CHUNG VE TRAO OI CHAT

C s cua s song la qua trnh bien hoa vat chat xay ra lien tuc trong te bao. Qua
trnh o goi la trao oi chat. Danh t trao oi chat am ch toan bo he thong cac phan
ng co tnh nh hng va lien quan mat thiet vi nhau xay ra trong te bao song nham
thc hien 4 chc nang ac trng sau ay:
1. Tiep nhan nang lng t moi trng dang nang lng hoa hoc cua cac
hp chat hu c hoac nang lng anh sang;
2. Bien hoa cac hp chat ngoai lai thanh vat lieu xay dng cua c the,
tc tien than cua cac ai phan t cua te bao;
3. Tong hp cac thanh phan protein, acid nucleic, lipid, polysaccharide va
cac thanh phan khac cua te bao t cac vat lieu xay dng noi tren;
4. Tong hp va phan huy nhng hp chat sinh hoc can cho viec thc hien
cac chc nang ac trng khac nhau cua te bao.
I. ONG HOA VA D HOA.
Trao oi chat bao gom hai qua trnh ong hoa va d hoa. D hoa la qua trnh phan
giai cac phan t thc an tng oi ln chu yeu bang cac phan ng oxy hoa. Nguon chat
dinh dng tham gia trong qua trnh d hoa c thu nhan t moi trng hoac t cac
kho d tr cua c the. Trong qua trnh d hoa cac phan t ln (glucid, lipid, protein) b
phan giai thanh nhng phan t nho hn nh acid lactic, acid acetic, ure, NH
3
, CO
2

v.v, ong thi giai phong nang lng t do von c cha trong cau truc phc tap cua
cac phan t hu c. e tr thanh hu ch, so nang lng nay c s bo tch luy lai
trong cac lien ket phosphate cao nang cua ATP va cac hp chat tng t.
ong hoa la qua trnh tong hp cac thanh phan tng oi ln cua te bao nh
polysaccharide, lipid, protein, acid nucleic t nhng hp chat tien than n gian. V
qua trnh nay lam tang kch thc phan t va lam cho cau truc phc tap hn, hay noi
cach khac, lam giam entropy cua he thong, nen no can tieu dung nang lng. Nang
lng nay c cung cap bang cach phan giai cac lien ket cao nang cua ATP.
Ca ong hoa va d hoa eu c cau thanh t hai qua trnh xay ra ong thi va
lien quan mat thiet vi nhau, o la:
1. Trat t cac phan ng enzyme dan en s phan huy hoac hnh thanh bo khung
cua mot phan t sinh hoc. San pham trung gian hnh thanh trong qua trnh nay c
goi la chat trao oi va toan bo chuoi bien hoa o c goi chung la trao oi trung
gian.
2. S bien hoa nang lng i kem vi moi mot phan ng enzyme cua qua trnh
trao oi trung gian. mot so giai oan cua qua trnh d hoa nang lng hoa hoc cua

Hoa sinh hoc - 64 -
cac chat trao oi c tch luy (thng di dang cua cac lien ket phosphate cao
nang); nhng giai oan nhat nh cua qua trnh ong hoa nang lng nay c em
ra chi dung. Phng dien nay cua trao oi chat c quy c goi chung la s lien hp
nang lng.
Trao doI Irung gIan va IIen Iop nang Iuong Ia IaI qua IrnI IIen quan
nIau va pIu IIuoc nIau. V vay, kII ngIIen cuu Irao doI cIaI can pIaI
pIan IcI cIung moI cacI dong IIoI.
Qua IrnI d Ioa cac cIaI dInI duong cInI bao gom ba gIaI doan cIu
yeu sau day:
1/ PIan gIaI cac cIaI dInI duong IIanI cac don v cau Iruc:
poIysaccIasIde b pIan gIaI IIanI monosaccIarIde; IIpId IIanI gIycerIne va
acId beo; proIeIn IIanI amInoacId v.v
2/ So Iuong Ion cac IoaI Iop cIaI kIac nIau InI IIanI Irong gIaI
doan 1 duoc cIuyen Ioa IIanI moI so I cac IoaI Iop cIaI don gIan Ion. V
du gIycerIne va IaI ca cac IoaI monosaccIarIde deu b pIan gIaI IIanI
gIyceraIdeIyde--pIospIaIe va sau do IIanI aceIyI-coenzyme A; TaI ca 20
amInoacId deu b pIan gIaI IIanI moI vaI san pIam don gIan Ion nIu
aceIyI-coenzyme A, -ceIogIuIaraIe, suxInaIe, IumaraIe, oxaIoaceIaIe;
/ Cac san pIam InI IIanI o gIaI doan 2 b oxy Ioa IIanI cac san
pIam cuoI cung: CO
2
, H
2
O va NH
.
Qua IrnI d Ioa cung bao gom gIaI doan. Nguyen IIeu xay dung ban
dau Ia nIung san pIam InI IIanI o gIaI doan cua qua IrnI d Ioa. O
gIaI doan 2 cac Iop cIaI con Iuong doI don gIan Irong gIaI doan 1 duoc
pIuc Iap Ioa IIem moI buoc, de sang gIaI doan cIung duoc su dung Iam
nguyen IIeu cIo cac pIan ung sInI Iong Iop cac daI pIan Iu nIu
poIysaccIasIde, proIeIn, acId nucIeIc
Mac du dong Ioa va d Ioa Ia IaI qua IrnI IraI nguoc nIau nIung cac
san pIam Irung gIan cua IaI qua IrnI nay Irong nIIeu kIau kIong Irung
nIau. V du, qua IrnI pIan gIaI gIycogen IIanI acId IacIIc, nIu Ia se IIay
sau nay, duoc IIuc IIen nIo 12 enzyme; Irong kII do qua IrnI Iong Iop
gIycogen Iu acId IacIIc cI co 9 enzyme Ia cIung voI qua IrnI pIan gIaI,
enzyme con IaI duoc IIay IIe bang nIung enzyme kIac.
Su Ion IaI IaI con duong kIac nIau cua dong Ioa va d Ioa Ia Ioan
Ioan can IIIeI, v con duong d Ioa ve maI nang Iuong kIong IIe duoc su
dung cIo con duong d Ioa.
Cac con duong dong Ioa va d Ioa IIuong xay ra Irong cac co quan Iu
kIac nIau cua Ie bao. V du oxy Ioa acId beo xay ra Irong Iy IIe, con sInI
Iong Iop acId beo duoc IIuc IIen Irong Ie bao cIaI. NIo dac dIem dnI v

Hoa sinh hoc - 65 -
kIac nIau nay ma cac con duong dong Ioa va d Ioa co IIe xay ra dong
IIoI va kIong pIu IIuoc nIau.
Su kIac nIau gIua IaI con duong dong Ioa va d Ioa con IIe IIen o
cIo cIung duoc dIeu Ioa bang nIung co cIe kIong gIong nIau va doc Iap
nIau.
Tuy nIIen, IaI qua IrnI dong Ioa va d Ioa co nIung gIaI doan cIung
IIuong duoc goI Ia nIung con duong trung tom ma moI Irong nIung v du
dIen InI Ia cIu IrInI Krebs. Trong cIu IrnI nay, moI maI, cac Iop cIaI
Iuu co se b pIan gIaI den cung IIanI CO
2
, maI kIac, cIung cung co IIe
duoc su dung Iam nguyen IIeu cIo qua IrnI dong Ioa.
II. CAC HNH THC VAN CHUYEN NANG LNG TRONG TRAO
OI CHAT.
PIan Iu cua cac Iop cIaI Iuu co pIuc Iap, v du gIucose, do co muc
do Io cIuc cao nen cIua Irong no moI nang Iuong IIem Iang raI Ion, dong
IIoI enIropy cua no Iuong doI IIap. KII pIan Iu gIucose b pIan gIaI IIanI
6 pIan Iu CO
2
va 6 pIan Iu H
2
O, do muc do Io cIuc cua cac san pIam
IIap Ion nen enIropy cua Ie IIong Iang Ien voI muc do Iuong ung, dong
IIoI, moI pIan nang Iuong nIaI dnI gIaI pIong Irong qua IrnI nay duoc
su dung de IIuc IIen moI cong nao do. PIan nang Iuong nay duoc goI Ia
Nong tuong tu do.
Trong cac Ie IIong sInI Ioc nang Iuong Iu do bao gIo cung Ia Ioa
nang, v cI co Ioa nang moI duoc co IIe su dung de IIuc IIen cong do co
IIe Ia nIung Ie IIong dang nIIeI.

Hnh VIII.1. Coc hnh thuc uon chugen nong luong :

Hoa sinh hoc - 66 -
A Long chu trnh ADP ATP
R Long dten tu
Nang Iuong Iu do gIaI pIong Irong cac pIan ung d Ioa duoc so bo IcI
Iuy Irong cac IIen keI Ioa Ioc dac bIeI goI Ia tten ket coo nong cua ATP.
NIung pIan Iu ATP nay sau do dI vao nIung kIu vuc cua Ie bao ma o do
can dung IoI nang Iuong. TaI day no cIuyen moI Ioac IaI goc pIospIaIe
Ian cung cua mnI cIo moI pIan Iu cIaI nIan nao do va bang cacI ay
Iruyen nang Iuong Ioa Ioc von IcI Iuy duoc Iruoc do cIo pIan Iu nay, Iam
cIo no co kIa nang IIuc IIen cong. Han IIan ATP sau kII maI cac goc
pIospIaIe Ian cung se bIen IIanI ADP Ioac AMP de roI IaI keI Iop IIem
cac goc pIospIaIe moI Irong cac pIan ung IIen Iop voI cac pIan ung gIaI
pIong nang Iuong de Iao nen nIung pIan Iu ATP moI. NIu vay, nang Iuong
Iu do Irong Ie bao duoc van cIuyen o dang cac goc pIospIaIe Ian cung
cua ATP nIo kIa nang cIuyen Ioa IIuan ngIcI ATP ADP.
NgoaI cIu IrnI ATP ADP noI Iren nang Iuong con duoc van
cIuyen Irong Ie bao o dang cac dIen Iu (xem cIuong Trao doI gIucId). e
Iong Iop cac Iop cIaI Iuu co Irong Ie bao IIuong pIaI cung cap dIen Iu va
H
+
. NIung dIen Iu nay duoc cIuyen den cac cIaI nIan Iu cac cIaI cIo
Irong cac pIan ung oxy Ioa kIu voI su IIam gIa cua moI so coenzyme
dac bIeI cIuyen Iam nIIem vu van cIuyen dIen Iu. Quan Irong nIaI Irong
so cac coenzyme IoaI nay Ia NADP. No dong vaI Iro nIu moI cIaI mang cac
dIen Iu gIau nang Iuong Iu cac san pIam d Ioa den cac pIan ung dong
Ioa von can duoc cung cap nang Iuong o dang nay. Co IIe xem vaI Iro cua
NADP Irong van cIuyen dIen Iu Iuong Iu nIu vaI Iro cua ATP Irong van
cIuyen cac goc pIospIaIe gIau nang Iuong.
Trong Ie bao IaI kIeu van cIuyen nang Iuong nay IIen quan maI IIIeI
voI nIau: nang Iuong Iu do gIaI pIong kII van cIuyen dIen Iu Iu moI Ie
IIong co nang Iuong cao den moI Ie IIong co muc nang Iuong IIap duoc so
bo IcI Iuy Irong cac pIan Iu ATP Iruoc kII dung de IIuc IIen cong.
III. NANG LNG SINH HOC VA CHU TRNH ATP
HIen IIIen nang Iuong Irong cac pIan ung sInI Ioa Ioan Ioan Iuan
IIeo cac dnI IuaI cua nIIeI dong Ioc.
Hen canI dnI IuaI IIu nIaI ve bao Ioan nang Iuong, dnI IuaI IIu IaI
cua nIIeI dong Ioc quy dnI cIIeu Iuong bIen Ioa Iu pIaI cua nang Iuong
Irong kII xay ra cac bIen doI vaI Iy Ioac cac pIan ung Ioa Ioc. TIeo dnI
IuaI nay moI qua IrnI deu co xu Iuong xay ra IIeo cIIeu Iam Iang enIropy.
Xu Iuong nay Ion IaI cIo den kII daI duoc Irang IIaI can bang ma o do
enIropy co gIa Ir IoI da Iuong ung voI nIIeI do va ap suaI nIaI dnI.
EnIropy duoc xem Ia IIeu cIuan danI gIa muc do Ion Ioan cua Ie IIong,
con Irang IIaI can bang duoc xem Ia Irang IIaI ma IaI do kIong con xay ra

Hoa sinh hoc - 67 -
cac bIen doI vaI Iy va Ioa Ioc cua Ie IIong; Ie IIong o Irang IIaI can
bang kIong con kIa nang IIuc IIen cong. Sau kII Irang IIaI can bang,
cung voI vIec Iang enIropy no kIong con kIa nang Iu IIay doI cIIeu de
quay Iro IaI Irang IIaI ban dau v nIu vay co ngIIa Ia gIam enIropy. Cac
qua IrnI kIong Iam bIen doI enIropy duoc goI Ia guo trnh thuon ngh|ch,
con cac qua IrnI Iam Iang enIropy duoc goI Ia guo trnh khong thuon
ngh|ch.
EnIropy Ia moI Iam so Ioan Ioc duoc xac dnI boI nIIeu IIong so,
Irong do co nIIeI do va ap suaI. on v do enIropy Ia caIo/
o
C.
HIen IIIen enIropy Irong cac pIan ung Ioa Ioc IIen quan voI bIen
IIIen nang Iuong Iu do. MoI IIen Ie nay Irong cac Ie IIong sInI Ioc duoc
xac dnI bang pIuong IrnI:
E = O + TS
Irong do E Ia bIen IIIen Iong nang Iuong cua Ie IIong; O Ia bIen
IIIen nang Iuong Iu do cua Ie IIong, S Ia bIen IIIen enIropy, T Ia nIIeI
do IuyeI doI.
HIen IIIen nang Iuong Iu do O Ia pIan bIen IIIen nang Iuong cua Ie
IIong co IIe duoc su dung de IIuc IIen cong Irong qua IrnI ma Ie IIong
Iuong ve Irang IIaI can bang Irong dIeu kIen nIIeI do va ap suaI kIong
doI. Trang IIaI can bang Ia Irang IIaI Iuong ung voI gIa Ir nIo nIaI cua
nang Iuong Iu do.
HIen IIIen nang Iuong Iu do IIuong duoc dung de xac dnI cIIeu
Iuong cua moI pIan ung Ioa Ioc. Trong dIeu kIen nIIeI do va ap suaI
nIaI dnI nang Iuong Iu do cua Ie IIong co xu Iuong gIam den gIa Ir IoI
IIIeu Iuong ung voI Irang IIaI can bang.
OIa Ir bIen IIIen nang Iuong Iu do O cua pIan ung Ioa Ioc nIo
pIuong IrnI ruI ra Iu dnI IuaI can bang Ioa Ioc.
oI voI pIan ung baI ky

A + bH cC + dD
(1)
Irong do a, b, c, va d Ia so pIan Iu Iuong ung cua cac cIaI A, H, C va D.
HIen IIIen nang Iuong Iu do cua pIan ung (1) bang:

C}
c
|D}
d

O =
O
o
+ RTIn (2)
|A}
a
|H}
b


Hoa sinh hoc - 68 -
Irong do |A}, |H}, |C} va |D} Ia nong do pIan Iu gam (moI) cua cac cIaI A, H,
C va D; R Ia Iang so kI bang 1,987 CaIo/moI.
o
C; T Ia nIIeI do IuyeI doI;
O
o
Ia bIen IIIen nang Iuong Iu do IIeu cIuan, Iuc bIen IIIen nang Iuong Iu
do cua pIan ung Ioa Ioc kII pH = 0, nong do cac cIaI IIam gIa pIan ung
va san pIam cua pIan ung deu bang 1.0M, ap suaI 1 aIm, nIIeI do
25
o
C.
oI voI pIan ung (1) IaI Irang IIaI can bang, kIong pIu IIuoc vao
nong do cua cac cIaI A, H, C va D, nang Iuong Iu do co gIa Ir IoI IIIeu va
no kIong IIep Iuc bIen IIIen nua, Iuc O = 0, co ngIIa Ia:

|C}
c
|D}
d

0 = O
o
+ RTIn

|A}
a
|H}
b
()

|C}
c
|D}
d

O
o
= - RTIn

|A}
a
|H}
b
(4)

aI Iang so can bang cua pIan ung Ia K
eq
, Ia co:

|C}
c
|D}
d

K
eq
=

|A}
a
|H}
b

(5)
Tu do: O
o
= - RTInK
eq
, Iay: O
o
= -
2,0RTIgK (6)
eq

Tu pIuong IrnI (6) Ia IIay, neu bIeI duoc Iang so can bang K
eq
cua
pIan ung o nIIeI do baI ky cIo Iruoc II Ia co IIe xac dnI duoc gIa Ir
bIen IIIen nang Iuong Iu do IIeu cIuan O
o
cua pIan ung do. NIu vay, O
o
Ia moI Iang so nIIeI dong Ioc dac Irung cIo moI pIan ung Ioa Ioc. Qua
pIuong IrnI (2) Ia co IIe IIay rang O
o
se bang O kII nong do cua moI
cIaI IIam gIa pIan ung deu bang 1,0M.

Hoa sinh hoc - 69 -
HIen IIIen nang Iuong Iu do IIeu cIuan O
o
cua pIan ung cung co IIe
duoc xem Ia IIeu so gIua nang Iuong Iu do IIeu cIuan cua san pIam cua
pIan ung O
o
p
va nang Iuong Iu do IIeu cIuan O
o
r
cua cac cIaI ban dau,
Iuc:

O
o
= O
o
r
- O
o
r

(7)
O
o

doI voI moI pIan ung IaI moI nIIeI do nIaI dnI Ia moI Iang so,
con O cua pIan ung bIen doI Iuy IIuoc vao nong do cua cac cIaI ban dau
va san pIam cua pIan ung. OIa Ir O cIo IIay pIan ung co IIe xay ra
Iay kIong o dIeu kIen nong do nIaI dnI cua cac cIaI IIam gIa pIan ung.
PIan ung cI co IIe xay ra Irong Iruong Iop O co gIa Ir am, Iuc Irong
Iruong Iop nang Iuong Iu do gIam. Can Iuu y rang moI pIan ung voI O
o
>0
van co IIe xay ra, mIen Ia I Ie nong do cua san pIam cua pIan ung va
cua cac cIaI ban dau Iam cIo O< 0.
PIuong IrnI (6) cIo pIep InI O
o
cua baI ky pIan ung Ioa Ioc nao
Iren co so gIa Ir K
eq
. Neu K
eq
= 1 II O
o
= 0; neu K
eq
> 1 II O
o
< 0,
pIan ung Irong Iruong Iop do Ia pIan ung gIaI pIong nang Iuong; neu K
eq

< 1 II O
o
> 0 va pIan ung Irong Iruong Iop do Ia pIan ung IIu nIan
nang Iuong. LoaI pIan ung nay se kIong Iu xay ra IIeo cIIeu IIuan neu
nong do ban dau cua cac cIaI IIam gIa pIan ung bang 1,0M.
TaI ca nIung dIeu neu Iren deu dung voI cac pIan ung sInI Ioa. Tuy
nIIen, kII pIan IcI nIIeI dong Ioc cua cac Ie IIong sInI Ioa can pIaI bo
sung ba dIeu kIen quan Irong sau day:
1. Neu cIaI pIan ung ban dau Ioac san pIam cua pIan ung Ia
nuoc II nong do cua no Iuon duoc xem Ia bang 1,0M;
2. OIa Ir pH cIuan cua cac Ie IIong sInI Ioa duoc xem Ia pH =
7 cIu kIong pIaI pH = 0, v vay bIen IIIen nang Iuong Iu do IIeu cIuan cua
cac Ie IIong sInI Ioa duoc ky IIeu Ia O
o
;
. Su dung gIa Ir O
o
Irong vIec xac dnI bIen IIIen nang Iuong
Iu do IIeu cIuan cua cac Ie IIong sInI Ioa co ngIIa Ia cac Irang IIaI Ion
Ioa va kIong Ion Ioa o pH = 7 cua cac cIaI IIam gIa pIan ung va san
pIam cua pIan ung Ia Irang IIaI cIuan. KII pH IIay doI, muc do Ion Ioa
cua moI Ioac moI so IIanI pIan cung bIen doI IIeo.
TIong qua vIec xac dnI Iang so can bang K
eq
nguoI Ia da xac dnI
duoc gIa Ir O
o
cua cac pIan ung IIuy pIan ATP sau day:
ATP + H
2
O
ADP + P
vc
: O
o
= -7,KcaI/moI
ADP + H
2
O
AMP + P
vc
: O
o
= -7,KcaI/moI

Hoa sinh hoc - 70 -
AMP + H
2
O
AdenosIne + P
vc
: O
o
= -,4KcaI/moI
Tren co so cac gIa Ir O
o
cua cac pIan ung IIuy pIan Iren day nguoI
Ia goI cac IIen keI cua IaI goc pIospIaIe Ian cung cua pIan Iu ATP Ia tten
ket phosphote coo nong. Tuy nIIen, neu so sanI O
o
cua cac IIen keI
pIospIaIe cao nang voI O
o
cua cac pIan ung IIuy pIan nIIeu Iop cIaI
kIac (bang VIII.1), Ia se IIay gIa Ir 7,KcaI cI cIIem v Ir Irung gIan.
CInI nIo dac dIem nay ma ATP co IIe dong vaI Iro quan Irong Irong vIec
van cIuyen cac goc pIospIaIe Iu cac IIen keI gIau nang Iuong den cac cIaI
ngIeo nang Iuong de Iam cIo nIung cIaI nay Iro nen gIau nang Iuong Ion.
Rong VIII.1. Gto trj G
o
cuo phon ung thug phon not so phosphote co g nghto
stnh hoc.
Co chot G
o
Co chot G
o

PIospIoenoIpyruvaIe - 14,80 ATP -7,0
1,-dIpIospIogIyceraIe -11,80 OIucoso-1-pIospIaIe - 5,00
CreaIIne pIospIaIe -10,0 FrucIoso-6-pIospIaIe - ,8
AceIyIpIospIaIe -10,10 OIucozo-6-pIospIaIe - ,
ArgInInepIospIaIe -7,70 OIyceroIpIospIaIe -2,00
Trong cac pIan ung noI IIep nIau ma Irong do vIec van cIuyen nang
Iuong do ATP IIuc IIen, nang Iuong Ioa Ioc Iruoc IIen duoc Iruyen Iu moI
Iop cIaI gIau nang Iuong, v du pIospIoenoIpyruvaIe Ioac 1,-
dIpIospIogIyceraIe (co O
o
> 7,KcaI) sang ADP de duoc Iam IIoI IcI Iuy
Irong moI Ioac IaI IIen keI cao nang cua ATP. Sau do ATP IaI nIuong moI
Ioac IaI goc pIospIaIe Ian cung cua mnI cIo pIan Iu cIaI nIan von
ngIeo nang Iuong, Iam cIo suc cIua nang Iuong, va do do IoaI InI Ioa
Ioc, cua no duoc Iang Ien.
NgoaI ATP, nang Iuong con co IIe duoc van cIuyen nIo cac nucIeosIde
IrIpIospIaIe kIac nIu OTP, UTP v.v... Tuy nIIen, su pIuc IoI cac
nucIeosIde IrIpIospIaIe nay van pIaI nIo den ATP voI su xuc Iac cua
enzyme nucIeozIdedIpIospIaIe kInase:
ATP + UDP ADP + UTP
ATP + CDP ADP + CTP
ATP + ODP ADP + OTP
IV. VAN CHUYEN NANG LNG TRONG CAC PHAN NG OXY
HOA KH
PIan ung oxy Ioa kIu Ia pIan ung ma Irong do dIen Iu duoc cIuyen
Iu cIaI cIo (cIaI kIu) sang cIaI nIan (cIaI oxy Ioa) (InI VIII.1H). Trong
co IIe song qua IrnI nay duoc IIuc IIen nIo su IIam gIa cua cac enzyme
oxy Ioa kIu kIac nIau ma coenzyme cua cIung dong vaI Iro nIu nIung

Hoa sinh hoc - 71 -
cIaI van cIuyen dIen Iu. Truoc IIen, dIen Iu duoc IacI ra Iu cIaI cIo de
keI Iop voI moI coenzyme nao do o dang oxy Ioa (NAD
+
, NADP
+
, FAD...) de
cIuyen nIung coenzyme nay sang dang kIu (NAD.H, NADP.H, FAD.H
2
...);
sau do Iu nIung coenzyme o dang kIu nay dIen Iu duoc cIuyen den cIo
moI cIaI nIan co muc nang Iuong IIap Ion.
CIaI oxy Ioa va cIaI kIu bao gIo cung IoaI dong IIen Iop voI nIau
Iung cap. KIa nang xay ra pIan ung oxy Ioa kIu duoc danI gIa Iren co
so the khu E, Iuc IIeu so dIen IIe (do bang voII) gIua cap cIaI kIu cIaI
oxy Ioa Irong pIan ung.
TIe kIu cua moI pIan ung oxy Ioa kIu xay ra Irong dIeu kIen ap
suaI bang 1,0 aIm, nIIeI do bang 25
o
C, pH bang 0, nong do cIaI oxy Ioa
va cIaI kIu deu bang 1.0 moI duoc goI Ia the khu tteu chuon, ky IIeu Ia
E
o
, cua pIan ung do.
TIe kIu cua pIan ung H
2
2H
+
+ 2e
-
, xay ra Irong dIeu kIen noI
Iren duoc xem Ia don v cua IIe kIu IIeu cIuan (E
o
= 1,0).
Do Irong cac Ie IIong sInI Ioc dIeu kIen IIuan IoI nIaI de Iau IeI
cac pIan ung sInI Ioa xay ra Ia o pH = 7 nen IIe kIu IIeu cIuan cua cac
pIan ung sInI Ioa duoc quy uoc xac dnI o pH = 7 va ky IIeu Ia E
o
. Neu
E
o
cua pIan ung H
2
2H
+
+ 2e
-
bang 1,0 II E
o
cua no bang 0,42V.
TIe kIu IIeu cIuan cua moI so cap cIaI oxy Ioa cIaI kIu quan
Irong duoc gIoI IIIeu Irong bang VIII.2.
Cac Ie IIong
co gIa Ir am cua
IIe kIu IIeu cIuan
cao Ion so voI cap
H
2
2H
+
se co kIa
nang nIuong dIen
Iu Ion Ion kIa
nang nay cua
Iydro; nguoc IaI,
cac Ie IIong co gIa
Ir duong cua IIe
kIu IIeu cIuan Ion
Ion (Iay gIa Ir am
nIo Ion) II kIa
nang nIuong dIen
Iu cua cIung IIap
Ion so voI Iydro.
Bang VIII.2. The kh tieu chuan cua mot so he thong oxy
hoa kh sinh hoc.
Chot khu Chot oxg hoo
o
(V)
AceIaIdeIyde AceIaIe -0,60
H
2
2H
+
-0,42
IsocIIraIe -
CeIogIuIaraIe
-0,8
NAD.H+H
+
(NADP.H+H
+
) NAD
+
(NADP
+
) -0,2
FAD.H
2
FAD -0,11
CyIocIrome b (Fe
2+
) CyIocIrome b
(Fe
+
)
0,00
CyIocIrome c (Fe
2+
) CyIocIrome c +0 26
TIe kIu IIeu cIuan E
o
va IIe kIu bIeu kIen E
I
cua moI pIan ung oxy
Ioa kIu IIen Ie voI nIau bang pIuong IrnI:

Hoa sinh hoc - 72 -

2,0RT |CIaI nIan e
-
}
E
I
= E
o
+ Ig

nF |CIaI cIo e
-
}
Irong do R Ia Iang so kI (8,1 |un/moI.do), T Ia nIIeI do IuyeI doI, n Ia so
dIen Iu duoc van cIuyen, F Ia so Faraday (96,406 |un/VoII).
V Irong cac Ie IIong sInI Ioa pIo bIen co cIe van cIuyen 2 dIen Iu
(n = 2), nen pIuong IrnI Iren co IIe duoc vIeI duoI dang:

|CIaI nIan
e
-
}
E
I
= E
o
+ 0,01 Ig

|CIaI cIo e
-
}
HIeI gIa Ir IIe kIu IIeu cIuan cua cac Ie IIong oxy Ioa kIu sInI
Ioc kIac nIau cIo pIep IIen doan cIIeu Iuong cua dong dIen Iu Irong cac
pIan ung oxy Ioa kIu Irong dIeu kIen IIeu cIuan IIeo nguyen Iac dong
dIen Iu cI co IIe cIuyen dong Iu pIaI Iu cIaI co gIa Ir am cua E
o
cao
den cIaI co gIa Ir am cua E
o
IIap Ion.
KII moI pIan ung oxy Ioa kIu xay ra cung voI su dI cIuyen cua
dIen Iu, nang Iuong Iu do cua Ie IIong cung bIen doI. MoI IIen Ie gIua bIen
IIIen IIe kIu IIeu cIuan (E
o
) va bIen IIIen nang Iuong Iu do IIeu cIuan
cua pIan ung (O
o
) duoc xac dnI bang cong IIuc:
O
o
= -nFE
o

Irong do n Ia so dIen Iu duoc van cIuyen, F Ia so Faraday
(2,062KcaI/VoII); cac IIong so duoc xac dnI Irong dIeu kIen IIeu cIuan,
Iuc IaI ca cac IIanI pIan deu co nong do 1,0 moI, nIIeI do 25
o
C, pH=7,0.
NIo pIuong IrnI Iren day co IIe InI bIen IIIen nang Iuong Iu do
IIeu cIuan cIo cac Ie IIong oxy Ioa kIu kIac nIau. V du, kII cap dIen
Iu (Ioac Iydro) duoc van cIuyen Iu NAD.H (E
o
= -0,2V) den oxy pIan Iu
(E
o
= +0,82V):

O
o
= -2 x 2,062 x |0,82 (-0,2)} = -52,7 KcaI.
Nang Iuong Iu do gIaI pIong Irong cac pIan ung oxy Ioa kIu se co
IIe duoc dung de Iong Iop ATP Iu ADP va pIospIaIe vo co neu IIoa man
IaI dIeu kIen: 1/ Ion IaI Ie enzyme pIospIoryI Ioa IIen Iop voI Ie enzyme
van cIuyen dIen Iu; 2/ bIen IIIen nang Iuong Iu do du Ion (>7,KcaI/moI).


Hoa sinh hoc - 73 -
CHNG 4. GLUCID

Glucid la nhom hp chat hu c pho bien rong rai trong t nhien ma ban chat
hoa hoc la dan xuat aldehyde hoac cetone cua ru a chc (polyalcohol) hoac la san
pham ngng tu cua nhng dan xuat nay. V t le gia H va O trong nhieu loai glucid
giong nh t le gia nhng nguyen to nay trong nc, va ben canh o con co nguyen to
carbon, nen trc ay ngi ta thng goi nhom hp chat nay la hydrate carbon. Tuy
nhien, cach goi nay ngay nay t c dung. o la do ngi ta phat hien c ngay
cang nhieu loai glucid co t le gia H va O khong giong nh trong nc; hn the na,
ngi ta cung a tm thay mot so loai glucid ma phan t cua chung co cha nit (v du
glucosamine, galactosamine v.v...).
Y ngha cua glucid oi vi i song cua sinh vat la vo cung to ln. thc vat
glucid chiem 25-90% chat kho. Chung la san pham chu yeu cua quang hp va c
tch luy trong cac c quan khac nhau cua cay e lam chat dinh dng d tr. Mot so
glucid lam nhiem vu nang va gop phan chu yeu vao viec kien tao vach te bao thc
vat. ong vat ham lng glucid thng khong vt qua 2% chat kho, tch luy chu
yeu trong gan va c dang hp chat cao phan t glycogen. Tuy nhien ieu o khong
co nghia la glucid t can thiet oi vi i song ong vat, bi v phan ln nang lng
can cho qua trnh hoat ong song cua ong vat cung nh cua thc vat la do glucid
cung cap.
Cac san pham chuyen hoa trung gian cua glucid trong c the song la nguyen lieu
e tong hp nhieu loai hp chat khac nhau. Chung con la thanh phan cau tao cua
nhieu loai hp chat cc ky quan trong nh acid nucleic, mot so coenzyme, cac hp
chat cao nang, cac hp chat quy nh nhom mau v.v...
Glucid c chia lam hai nhom ln: monosacharide (monose) va polysacharide
(polyose). Phan t polysacharide cha t hai goc monose tr len. Nhng
polysacharide cha so goc monose khong nhieu lam trong phan t con c goi la
oligosaccharide (disacharide, trisacharide, tetrasaccharide v.v...). Cung nh
monosacharide, oligo-saccharide de tan trong nc, che pham tinh khiet co dang tinh
the, co v ngot, do o c goi chung la ng. Tuy nhien, nhng oligosaccharide co
phan t tng oi ln khong tan trong ethanol nh monosacharide va cac
oligosaccharide phan t nho. Cac polysacharide co phan t ln khong tan trong nc
(celluluse) hay tao trong nc nhng dung dch keo rat nht (tinh bot, glycogen,
hemixellulose, pectin, chat nhay v.v...).
I. MONOSACHARIDE (MONOSE)
1. cau tao.
Monosacharide la nhng polyoxyaldehyde hoac polyoxycetone cua mot so ru
a chc (polyalcohol). Mot trong nhng ru a cha n gian nhat la glycerine

Hoa sinh hoc - 74 -

(glycerol). Khi chc ru bac 1 cua no b oxy hoa se tao ra aldehyde glyceric
(glyceraldehyde); neu chc ru bac 2 b oxy hoa se tao ra dioxyacetone. Nhng
monose nay cha 3 nguyen t carbon nen c goi chung la triose. Nhng
monosacharide cha 4, 5, 6, 7 nguyen t carbon co ten goi tng ng la tetrose,
pentose, hesose va heptose.
Mat khac, phu thuoc vao ac iem phan t
monose cha nhom aldehyde hay cetone ma
chung c xep vao nhom aldose hay cetose.
Glyceral-dehyde thuoc nhom aldose (aldotriose),
con dioxyacetone thuoc nhom cetose (cetotriose).
Cung nh aminoacid, trong phan t monose,
tr dioxyacetone, co cha mot hay nhieu nguyen
t carbon bat oi, nen chung co the ton tai cac
dang ong phan quang hoc D- hoac L- vi hoat
tnh quang hoc ac trng. So lng ong phan
quang hoc cua moi loai monose c xac nh
bang cong thc X = 2
n
, trong o n la so nguyen t
carbon bat oi.
D- va L- glyceraldehyde co cau tao nh mo
ta trong hnh ben canh.
oi vi cac monose khac neu cac nhom chc gan vi nguyen t carbon bat oi
xa chc aldehyde hoac cetone nhat co s phan bo trong khong gian giong D-
glyceraldehyde th c xep vao nhom D-; neu giong L-glyceraldehyde th c xep
vao nhom L-.V du, D-glucose va L-glucose co cau tao nh sau:
Trong t nhien monose thng ton tai
dang D. Ngoai D-glyceraldehyde va D-
glucose, hang loat cac monose khac cung
ong vai tro rat quan trong trong cac qua trnh
hoat ong song. o la D- erytrose, D-ribose,
D-arabinose, D-xylose, D-galactose, D-
mannose (thuoc nhom aldose) va D-fructose,
D-ribulose, D-xylulose, D-cedoheptulose, D-
mannoheptulose (thuoc nhom cetose) cung
nh mot so ng deoxy (2-D-deoxyribose, L-
rhamnose, L-fucose) va ng amin (D-
glucosamine, D-galactosamine)

Hoa sinh hoc - 75 -

Khac vi aminoacid va nhieu loai hp chat co hoat tnh quang hoc khac, tat ca
cac monose cha t 4 nguyen t carbon tr len khi tan trong nc se thay oi gia tr
hoat tnh quang hoc. Hien tng nay c goi la s chuyen quay (mutarotation).
Nguyen nhan cua hien tng nay la cho ben canh cau truc mach h nh a gii
thieu tren, nhng monose nay con co kha nang ton tai dang cau truc vong. Nhng
dang mach vong nay do lam xuat hien them mot nguyen t carbon bat oi nen co hoat
tnh quang hoc khac vi dang mach h. Gia tr goc quay cua dung dnh la gia tr trung
bnh cua cac dang cau truc o. V du, D-glucose trong dung dch co them hai dang cau
truc mach vong la --D-glucose va -D-glucose:
Trong so ba dang cau truc
nay dang mach h chiem t le
khong ang ke, dang -D- chiem
khoang 1/3 vi o quay rieng bang
+112,2
o
, con dang -D- chiem
khoang 2/3 vi o quay rieng bang
+18,7
o
. T le nay quyet nh o
quay rieng cua dung dch D-
lgucose
trong nc la +52,7
o
.

Hoa sinh hoc - 76 -
S hnh thanh cau truc vong la ket qua cua phan ng tao semiacetal noi. oi vi
aldose phan ng nay xay ra nh sau:
O OH
R C + HO CH
2
R R C O CH
2
R
H H
hoac oi vi cetose:
O OH
R C + HO CH
2
R R C O CH
2
R
CH
2
OH CH
2
OH

Qua trnh nay dan en s xuat hien them mot nguyen t carbon bat oi (C
1
oi
vi aldose va C
2
oi vi cetose) va mot nhom OH (gan vi carbon bat oi o). Nhom
OH nay c goi la nhom hyrdoxyl semiacetal. No co hoat tnh hoa hoc cao hn
nhieu so vi cac nhom OH khac. Tuy thuoc vao v tr khong gian cua nhom OH
semiacetal ma cau truc vong cua monose co dang - hay dang -. Mot monose mach
vong se thuoc dang - neu nhom OH semiacetal co v tr khong gian cung pha vi
nhom OH gan vi nguyen t carbon bat oi von quyet nh monose o thuoc day D-
hay day L-. Trong trng hp ngc lai monose mach vong se thuoc dang -.
Cac dang mach vong tren ay cua glucose va nhng dang vong tng t co the
c xem la dan xuat cua pyran va v the chung c xep vao nhom pyranose
(glucopyranose, galactopyranose v.v...).
Cau truc vong cua monose con co the c hnh thanh dang vong 5 canh (cau
oxy noi cac nguyen t C
1
vi C
4
aldose hoac C
2
vi C
5
cetose). Cau truc nay co
the c xem la dan xuat cua furan nen c goi chung la furanose.

e mo ta cau truc khong gian cua monose vong ngi ta s dung mot kieu cong
thc co ten la cong thc phoi canh. Theo cach dien at nay cac nguyen t C
1
C
5


Hoa sinh hoc - 77 -
cua glucopyranose hoac C
2
C
5
cua fructofuranose cung vi nguyen t oxy lam thanh
mat phang nam ngang vi net am hng ve pha trc, tc pha ngi nhn; cac nhom
chc gan vi cac nguyen t carbon nay (-H, -OH, CH
2
OH...) c phan bo pha tren
hoac pha di mat phang o. Quan sat v tr cua nhom OH semiacetal trong cong
thc phoi canh, ta co the phan biet c de dang cac dang - va - cua mot monose
vong. Tuy nhien cach dien at nay co the gay an tng sai lam rang vong pyran hay
vong furan co cau truc phang. Tren thc te cac cau truc vong pyranose co the co cau
truc dang ghe hay dang thuyen, trong o dang ghe ben vng hn. Ngi ta cho rang
cac loai ng hexose trong t nhien ton tai dang nay. Cac nhom chc gan vi vong
pyran dang thuyen hay dang ghe c chia lam hai nhom: nhom truc (a) va nhom
xch ao (e), trong o cac nhom OH xch ao de tham gia cac phan ng esetr hoa
hn cac nhom OH truc.

2. Tnh chat hoa hoc.
a/ Phan ng tong hp glycoside: Thong qua nhom OH semiacetal von co hoat
tnh hoa hoc cao cac monose co the ket hp vi nhieu loai hp chat khac nhau e tao
nen cac san pham co ten chung la glycoside. Tuy thuoc nhom OH semiacetal co v
tr - hay - ma glycoside c chia lam hai nhom: -glycoside va -glycodise vi
tnh chat rat khac nhau, ac biet la trong quan he vi enzyme.
V du n gian nhat cua glycoside la - va -methylglycoside.


Hoa sinh hoc - 78 -
Do nhom OH semiacetal tham gia trc tiep trong viec tao thanh cac glycoside
nen no con c goi la nhom hydroxyl glycoside. Bo phan khong phai glucid trong
phan t glycoside c goi la nhom aglycon. No co the la goc ru, cac hp chat vong
thm, vong thm hydrogen hoa, steroid, alcaloid v.v...
Glycoside pho bien rong rai trong t nhien, ac biet la trong gii thc vat. Tren
c s ac iem cua lien ket gia hai thanh phan glucid va aglycon ngi ta phan
biet O-glycoside (RCOA), S-glycoside (RCSA), N-glycoside (RCNA),
va C-glycoside (RCCA). Pho bien nhat la nhom O-glycoside (v du: glucovanilin,
amigdalin, cac loai glycoside tim v.v...) va N-glycoside (v du cac loai nucleoside).
Glucovanilin co nhieu trong qua vani (Vanilla); thanh phan aglycon cua no la
vaniline, mot chat thm quy gia. Amigdaline co trong hat m, tao, man, ieu...
Aglycon cua no la hp chat gia acid benzoic va acid cyanhydric. Do co cha nhom
CN nen amigdalin co the lam cho ngi va gia suc b trung oc do c che ho hap.
Glycoside tim la mot nhom glycoside ma aglycon la cac dan xuat khac nhau cua
cyclopentanoperhydro-phenantren. Chung co tac dung rat manh len c tim.
Nucleoside la thanh phan cau tao cua acid nucleic va cua nhieu hp chat sinh hoc
quan trong khac (coenzyme, hp chat cao nang v.v...).

b/ Phan ng ester hoa. Thong qua cac nhom OH t do cua mnh, ac biet la cac
nhom OH hai au tan cung, monose phan ng vi cac acid cha oxy e tao ra cac
ester. Quan trong nhat la cac ester phosphate. Nhng ester nay co hoat tnh hoa hoc
rat cao va de dang tham gia hang loat phan ng cua qua trnh trao oi chat.. V du
glucoso-6-phosphate, glucoso-1-phosphate, fructoso-6-phosphate... co vai tro quan
trong trong chuyen hoa tinh bot va glycogen, trong cac qua trnh quang hp va ho hap.

c/ Phan ng oxy hoa va tnh kh cua monose: Khi monose b oxy hoa, tuy thuoc
vao ieu kien moi trng, co the hnh thanh cac san pham khac nhau. Neu b oxy hoa

Hoa sinh hoc - 79 -
trong moi trng acid, v du vi s tham gia cua nc brom, chc aldehyde cua
monose b oxy hoa va san pham thu c co ten chung la acid aldonic. V du glucose
b oxy hoa thanh acid gluconic.
Khi aldose b oxy hoa manh,
v du di tac dung cua acid nitric
ac, ca chc aldehyde va chc
ru bac mot eu b oxy hoa. San
pham la acid dicarboxylic co ten
chung la acid aldaric. V du
glucose b oxy hoa thanh acid
glucaric, galactose thanh acid
slisic, mannose thanh acid mannosaccharic...
Trong nhng trng hp ac biet (nh di tac dung cua enzyme), monose ch b
oxy hoa tai chc ru bac mot. Trong trng hp nay san pham co ten chung la acid
uronic (acid glucuronic, acid galacturonic, acid mannuronic...).
Trong c the thc vat acid uronic ton tai dang lien ket trong thanh phan cua
cac chat pectin, mot so loai chat nhay va nhng poylysacchride phc tap khac co ten
chung la polyuronide. Acid uronic con la san pham trung gian trong qua trnh chuyen
hoa hexose thanh pentose.
Tnh chat cua monose b oxy hoa bi cac chat oxy hoa yeu, v du dung dch
kiem cua oxyde ong II, c ng dung trong viec nh tch va nh lng ng.
Trong khi monose b oxy hoa th Cu
2+
b kh thanh Cu
+
. ac iem nay cua monose
c goi la tnh kh. Tat ca monose va nhng oligosacchride con co nhom OH
semiacetal t do (phan ln la disaccharide) c ac trng bi tnh kh va v the c
xep vao nhom ng kh.
Mot trong nhng san pham oxy hoa cua glucose acid glucuronic trong cac
mo cua thc vat va gan cua a so ong vat, tr ngi, vn, chuot bach va mot so loai
ong vat khac, la chat tien than e tong hp acid L-ascorbic, tc vitamin C (xem
chng 5 noi ve vitamin) .
Phan ng kh: Ngc lai vi phan ng oxy hoa, khi b kh monose chuyen hoa
thanh cac ru a chc (polyalcohol) tng ng: glyceraldehyde va dioxyacetone b
kh thanh glycerine; D-glucose va D-fructose thanh D-sobit(ol); D-galactose
thanh D-dulcit(ol); D-mannose thanh D-mannit(ol) v.v... Cac loai ru a chc nay
pho bien rat rong rai trong rau, qua, nam va tao.

Hoa sinh hoc - 80 -
II. OLIGOSACCHARIDE.
Da vao so goc monose trong phan t, oligosaccharide c chia thanh
disaccharide. trisacchride, tetrasaccharide v.v... e tao nen phan t oligosaccharide
cung nh polysac-charide, cac goc monose noi vi nhau bang cac lien ket glycoside.
1.Disacchride.
Trong phan t disaccharide hai goc monose lien ket vi nhau nh nhom OH
glycoside cua monose nay vi nhom OH bat ky cua monose kia. ac iem lien ket
gia cac goc monose co y ngha rat quan trong oi vi tnh chat cua disaccharide. Neu
hai nhom OH glycoside lien ket vi nhau th phan t disaccharide khong co tnh kh.
v du ien hnh cho nhom disaccharide nay la saccharose (trong cay ma, cay cu cai
ng) va trehalose (trong nam, tao, mot t thc vat bac cao va ong vat khong xng
song).

Neu trong so hai nhom OH tham gia tao thanh phan t disaccharide ch co mot
nhom OH glycoside th trong phan t disaccharide o con lai mot nhom OH
glycoside t do, lam cho phan t co tnh kh, va do o nhng polysacharide nay cung
vi monose c xep vao nhom ng kh. Pho bien nhat trong nhom disaccharide
nay la maltose, lactose, cellobiose, melibiose va gentiobiose.

Maltose la c s cau truc cua tinh bot va
glycogen; lactose cha trong sa ong vat, trong
ong phan cua mot so thc vat bac cao; cellobiose
la n v cau truc cua cellulose; melibiose la thanh

Hoa sinh hoc - 81 -
phan cau tao cua trisaccharide rhafinose;
gentiobiose la thanh phan
cau tao cua amygdaline va nhieu glycoside khac. Tat ca disaccharide di tac dung
cua acid hoac cua cac enzyme tng ng se b thuy phan thanh monose.
2.Trisaccharide.
ai dien cho nhom
trisaccharide la rhafinose.
No c phat hien trong
nhieu thc vat, ac biet la
trong hat bong va cu cai
ng. No khong co tnh
kh do ca 3
nhom OH glycoside eu
khong con trang thai t
do.

3.Tetrasaccharide.
ai dien cho nhom tetrasaccharide la stachiose. No c cau tao bi hai goc -
D-galactose, mot goc -Dglucose va mot goc -D-fructose. Co the xem phan t
stachiose la phan t rafinose c gan them mot goc -D-galactose th hai bang lien
ket glycoside 1-6 thong qua goc -galactose th nhat. Loai tetrasaccharide nay c
phat hien trong re va cu cua mot so thc vat va trong hat cay ho au. Cung nh
rafinose, stachiose khong co tnh kh.

III. POLYSACCHARIDE (POLYOSE).
Polyose hay polysacharide bac hai co trong lng phan t rat ln, hnh thanh t
rat nhieu goc monose. Tuy thuoc kch thc va ac iem cau truc cua phan t, chung
co the tao dung dch keo hoac hoan toan khong tan trong nc.

Hoa sinh hoc - 82 -
Nhng polysacharide hnh thanh t cung mot loai monose c goi la
homopolysac-charide; neu chung c tao nen t cac loai monose khac nhau th c
goi la hetero-polysaccharise.
1.Homopolisaccharide.
Trong t nhien phan ln cac homopolysaccharide la cac chat dinh dng d tr
(tinh bot, glycogen, dextran, inulin ...) hoac tham gia trong cau truc cua vach te bao
(cellulose, hemicellulose...)
Tinh bot la chat dinh dng chu yeu cua thc vat. Polysacharide tinh bot gom hai
loai co cau tao va tnh chat ly hoa hoc khac nhau: amylose va amylopectin.
Amylose de tan trong nc nong, tao nen dung dch keo khong nht lam. Dung
dch nay khong ben va khi e lang de b ket tua di dang tinh the. Amylopectin ch
tan trong nc un soi ap lc cao, tao nen dung dch keo rat nht va kha ben vng.
Trong lng phan t cua amylose vao khoang 300.000 1.000.000, con cua
amylopectin vai tram trieu. Amylose nhuom mau xanh vi iod, con amylopectin
mau o.
Trong phan t amylose cac goc -D-glucose noi vi nhau bang lien ket
glycoside 1-4, tao nen cau truc si khong phan nhanh, ton tai dang cau truc xoan oc,
moi vong xoan cha 6-7 goc glucose (hnh 2.1).

Hnh 2.1. S o cau truc xoan oc cua amylose
Trong phan t amylopectin ben canh lien ket glycoside 1-4 con co lien ket
glycoside 1-6 e tao ra cac mach nhanh vi cac iem phan nhanh cach nhau khoang
24 goc glucose.

Hoa sinh hoc - 83 -

Trong tinh bot cua cac loai thc vat khac nhau t le amylopectin / amylose khong
giong nhau. Trong bot gao t le nay vao khoang 17/83, con trong bot lua m 24/76.
Gia tr nay con phu thuoc vao giong, ieu kien canh tac va cac yeu to ngoai canh
khac.
Khi un vi acid, tinh bot b thuy phan thanh -D-glucose. Tinh bot hoa tan von
c s dung rong rai trong ky thuat phong th nghiem la san pham thuy phan khong
hoan toan cua tinh bot di tac dung cua acid loang.
Di tac dung cua enzyme amylase tinh bot b phan giai thanh maltose thong
qua cac san pham trung gian vi trong lng phan t nho dan co ten la dextrin.
-Amylodextrin nhuom mau xanh vi iod, tan trong ethanol 25% nhng b ket tua
bang ethanol 40% di dang bot trang;
- Erythrodextrin nhuom mau o vi iod, tan trong ethanol 55% nhng b ket tua
trong ethanol 65% di dang tinh the hnh cau;
- Achromodextrin khong nhuom mau vi iod, tan trong ethanol 70%;
- Maltodestrin co trong lng phan t nho nhat trong so cac destrin, khong
nhuom mau vi iod. Cung nh maltose va monosacharide, no hoa tan rat tot trong
ethanol 80%.
Glycogen, oi khi con c goi la tinh bot ong vat, co nhieu trong gan va c, la
nguon cung cap nang lng chu yeu cho moi qua trnh hoat ong song cua c the ong
vat. No co dang bot trang vo nh hnh, tan trong nc nong va tao thanh dung dch
keo mau trang uc. Khi tac dung vi iod, glycogen nhuom mau nau o hay tm o.
Trong lng phan t t 1 trieu (trong c) en 5 trieu (trong gan).
Cau tao cua phan t glycogen giong nh amylopectin nhng mc o phan nhanh
day hn. No b thuy phan di tac dung cua acid va enzyme giong nh tinh bot. Ngoai

Hoa sinh hoc - 84 -
ra, di tac dung cua phosphorylase va vi s tham gia cua phosphate vo c glycogen
b phan giai (theo mot c che enzyme co ten la phosphrolys) thanh glucoso-1-
phosphate.
Dextran la mot loai polysacharide ong vai tro chat dinh dng d tr cua vi
khuan va nam men. No c hnh thanh t cac goc -D-glucose noi vi nhau bang
cac lien ket glycoside 16. Cac mach nhanh c hnh thanh nh cac lien ket
glycoside 12, 1-3 hoac 14. Cac loai dextran khac nhau co mc o phan nhanh khac
nhau. Li dung ac iem nay ngi ta s dung dextran e che tao cac san pham co ten
la sephadex e s dung trong ky thuat phong th nghiem lam cac loai ray phan t.
Inulin la san pham quang hp va la chat dinh dng d tr cua mot so thc vat,
nh thc dc (Dahlia), diep xoan (Cicorium), actiso (artichaut) v.v... Phan t inulin
la mot mach dai khong phan nhanh c hnh thanh t 32 34 goc -D-fructose
thong qua lien ket glycoside 1 2. Do c hnh thanh t cac n v fructose nen
inulin c xep vao nhom polyfructoside.
Levan cung la mot loai polyfructoside. Khac vi inulin, trong phan t levan cac
goc fructose noi vi nhau bang cac lien ket fructoside 2 - 6. vi khuan cac nhom
OH t do trong phan t levan c metoxyl hoa. Levan cung co mat trong thc vat
thuoc ho Hoa thao, nhng cha t goc fructose hn va cac nhom OH khong b
metoxyl hoa.
Mot so polysacharide nhay tng t inulin va levan cung c cac vi khuan song
trong at tong hp nen va ong vai tro quan trong trong viec hnh thanh cau tng cua
at.
Trong so cac homopolysacharide la chat dinh dng d tr con co the ke en
xylan va mannan. Xylan hnh thanh t cac goc xylose, co mat trong cac mo thc vat.
Mannan hnh thanh t cac goc mannose. No la chat dung dch d tr cua vi khuan,
nam men, nam moc va thc vat bac cao.
Xylan, manan cung vi galactan (hnh thanh t cac goc galactose) va araban
(hnh thanh t cac goc arabinose) c goi chung la hemicellulose. Nhng
hemicellulose nay do c hnh thanh t mot loai monose duy nhat nen chung nam
trong nhom homopoly-saccharide. Ben canh chung con co nhng hemicellulose c
cau tao t mot so loai monose, va do o, theo nh ngha, chung thuoc nhom
heteropolysaccharide, Phan ln nhng hemicellulose nay tham gia trong cau truc cua
vach te bao cung vi cellulose.
Cellulose la thanh phan chu yeu cua vach te bao thc vat. n v cau truc cua
celllulose la -D-glucose. Chung noi vi nhau nh lien ket -1-4-glycoside, tao thanh
nhng mach dai khong phan nhanh. Trung bnh, moi phan t celllulose cha vai nghn
goc glucose. Cac si cellulose thng lien ket lai thanh bo khoang 60 phan t. S lien

Hoa sinh hoc - 85 -
ket nay c thc hien nh lien ket hydro gia cac nhom OH t do cua cac phan t
cellulose nam gan nhau.
Hnh 2.2. S o cau truc phan t cua amylose (A) va lien ket hydro gia cac si
cellulose nam gan nhau trong bo mach (B).
Cellulose khong tan trong nc, ru, eter nhng tan trong dung dch Cu(OH)
2

trong ammoniac dam ac (thuoc th Sweitzer). Acid sulfuric ac nhiet o soi thuy
phan cellulose thanh -D-glucose.
Hydro thuoc cac nhom OH t do trong phan t cellulose trong nhng ieu kien
nhat nh co the c thay the bang cac goc khac nhau (-CH
3
, CH
3
COO - v.v...) e tao
thanh cac dan xuat eter va ester. Nh cac phan ng nay t cellulose co the che tao
cellophan, celluloid, chat no, phim anh v.v... Nhieu dan xuat cua cellulose, nh
carboxycellulose (CM-cellulose), diethylaminoethyl-cellulose (DEAE-celllulose)
v.v... c s dung rong rai trong ky thuat phong th nghiem e phan oan protein,
acid nucleic bang phng phap sac ky trao oi ion.
2.Heteropolysaccharide.
Thuoc nhom heteropolysaccharide co nhieu loai hemicellulose, chat nhay va
gom, pectin, agar-agar, acid alginic, chitin, mucopolisaccharide...
Cac loai hemicellulose thuoc nhom heteropolysaccharide la nhng
polysacharide ma thanh phan cau tao cua chung gom cac loai acid uronic, arabinoa,
xylose va mot so monose khac. Nh a noi tren, chc nang chu yeu cua nhng
hemicellulose nay la tham gia trong cau truc cua vach te bao thc vat.
Chat nhay va gom la nhng polysacharide do thc vat tiet ra trong trang thai
sinh ly bnh thng (chat nhay) hoac khi bi ton thng (gom). Khi hoa tan trong nc
chung cho dung dch keo rat nht. Trong thanh phan cau tao cua hai loai
polysacharide nay co lactose, mannose, glucose, rhamnose, xylose va cac monose
khac.

Hoa sinh hoc - 86 -
Pectin la nhng polysacharide phan t ln, cha nhieu trong qua, cu va than cay.
Trong thc vat pectin ton tai dang khong tan protopectin. Sau khi s ly bang acid
loang, hoac di tac dung cua enzyme protopectinase, protopectin chuyen hoa thanh
pectin hoa tan. Qua trnh nay xay ra khi qua chn, lam cho qua tr nen mem.
Phan t pectin hoa tan
hnh thanh nh cac ester
methylic cua acid galacturonic
lien ket vi nhau bang lien ket
1-4-glycoside. Pectin t cac
nguon khac nhau co trong lng
phan t khong giong nhau, dao
ong t 20.000 en 50.000.
Pectin hoa tan trong nc b ket tua bang acetone 50%. Tnh chat ac trng cua
pectin la kha nang tao ra thach ong khi co mat acid va ng, do o no c s dung
rong rai trong ky nghe banh keo. Di tac dung cua kiem loang hoac cua enzyme
pectinase goc metoxyl (O CH
3
) b tach khoi chuoi polysaccharide va pectin b bien
thanh acid pectic (acid polygalacturonic), ong thi mat kha nang tao thach ong.
Agar-agar la mot loai polysacharide trong vach te bao cua mot so loai tao o
thuoc cac giong Gelidium, Gracilaria, Pterocladia va ahnfeltia. Agar-agar khong tan
trong nc lanh nhng tan trong nc nong di hnh thc dung dch keo. Dung dch
nay khi e nguoi ong lai thanh thach. Loai polysacharide nay khong c c the
ngi va ong vat hap thu. No c s dung rong rai trong y hoc va ky thuat phong th
nghiem trong viec lam moi trng nuoi cay vi sinh vat va nuoi cay mo thc vat.
Ngi ta cho rang agar-agar la hon hp cua t nhat hai loai polysacharide la
agarose va agaropectin. Agarose c cau tao bi cac goc D- va L-galactose noi vi
nhau bang lien ket 1-3-glycoside. Agaropectin hnh thanh t cac goc D-galactose va
mot so t goc galactoso- 6- sulfate. Tuy nhien, trong agar-agar con phat hien c cac
goc arabinose va glucose.
Acid alginic c phat hien trong vach te bao tao nau thuoc cac chi Macrocystis,
Laminaria, Fucus, Sargassum. o la mot loai polysacharide hnh thanh t cac goc acid
-D-mannuronic noi vi nhau bang lien ket 1-4-glycoside.
Acid alginic co kha nang tao dung dch keo nen c s dung rong rai trong
cong nghiep det e lam chat ho vai. No cung c dung e san xuat t nhan tao, lam
my pham...
Chitin la thanh phan chu yeu cua mo b cua con trung, tom, cua. No cung c
phat hien trong nam va a y. Trong cac mo ong vat chitin lien ket vi protein va
calcium carbonae. Phan t chitin rat giong cellulose nhng no c cau tao khong

Hoa sinh hoc - 87 -
phai t glucose ma t cac goc N-acetyl--D-glucosamine noi vi nhau bang cac lien
ket 1-4-glycoside.
Mucopolysaccharide ton tai trong cac mo ong vat nh sun, mo lien ket, trong
thanh phan cac chat gian bao, dch nhay... vi chc nang chu yeu la bao ve.
Thanh phan chu yeu cua mucopolysacchayride la glucosamine va acid uronic.
Trong cac mo ong vat chung ton tai mot phan trang thai t do, mot phan dang
lien ket vi protein (mucoprotein).
c hieu biet nhieu nhat trong so mucopolysacchayride la acid hyaluronic,
acid chondroitinsulfuric va heparin.
Acid hyaluronic co nhieu trong thuy tinh the cua mat, gan, dch khp, trong nang
cua mot so vi khuan va trong mang te bao trng. No c cau tao t N-acetyl--D-
glucosamine va acid D-glucuronic.

Di tac dung cua enzyme hyaluronidase do tinh trung tiet ra acid hyaluronic b
phan giai e tao ieu kien cho s thu tinh xay ra de dang. Cung nh acid hyaluronic,
cac khoang gian bao gi nc e te bao luon ton tai trong moi trng dung dch keo,
lam giam tac dung ma sat va chong lai s tham nhap cua vi trung.
Acid chondroitinsulfuric la thanh phan cua sun, xng, gan dang lien ket vi
protein collagen va lipid. Khi b thuy phan, acid chondroitinsulfuric se giai phong N-
acetylgalactosaminesulfate va acid galacturonic. Nhng goc nay cung noi vi nhau
bang cac lien ket -1-3 va 1-4-glycoside.
Heparin la
mot
heteropolysaccha
ride co tac dung
chong ong mau.
No c tong
hp va tch luy
trong gan. Ngoai
ra,


Hoa sinh hoc - 88 -
no con co trong phoi, tim, mat, tuyen giap trang, mau va trong nhieu c quan khac.
Tac dung chong ong mau cua heparin c thc hien nh no ngan can s chuyen hoa
prototrombin thanh trombin. Phan t heparin c cau tao t cac goc acid glucuronic
va -D-glucosamine dang dan xuat kep cua acid sulfuric.
IV. PHAN GIAI POLYSACCHARIDE.
1.Phan giai tinh bot va glycogen.
Moi polysaccharide trc
khi tham gia cac qua trnh trao
oi chat khac nhau eu can c
phan giai thanh monosaccharide.
Tinh bot va glycogen trong cac
mo thc vat va trong ng tieu
hoa cua ong vat c thuy phan
thanh maltose nh tac dung hp
ong cua ba enzyme: -amylase,
-amylase va -(1-6)-
glucosidase (hnh IX.1).
-Amylase (-1,4-glucan-
4-glu- canohydrolase) cat cac
lien ket (-1,4-glucoside, tao ra
san pham cuoi cung la hon hp
maltose va glucose thong qua san
pham trung gian la nhng
oligosacchaside cha 6-7 goc
glucose. Do khong the cong pha
lien
Hnh IX.1.Thuy phan amylopectin va glycogen
di tac dung cua -amylase, -amylase va -(1-6)-
glucosidase
ket -(1,6) nen -amylase cha lai nguyen ven khu vc phan nhanh cua amylopectin
va glycogen.
-Amylase (-1,4-glucan-4-glucanmaltohydrolase) phan giai amylose,
amylopectin va glycogen t nhng au tan cung khong kh cua phan t, tao ra san
pham cuoi cung la ng maltose. oi vi amylopectin va glycogen qua trnh dng lai
tai cac iem phan nhanh, e lai phan dextrin gii han.
-(1-6)-Glucosidase cong pha cac lien ket -(1-6)-glucoside. Nh o cac
dextrin gii han cha cac khu vc phan nhanh con lai sau tac dung cua - va -
amylase lai tiep tuc b thuy phan.
Khac vi qua trnh thuy phan trong ng tieu hoa, glycogen noi bao va tinh bot
mot so thc vat b phan giai bang con ng phosphorolis.

Hoa sinh hoc - 89 -
Trong qua trnh nay vi s tham gia cua acid phosphoric enzyme phosphorylase
tach goc glucose tan cung khong kh khoi phan t glycogen hoac tinh bot dang
glucoso-1-phosphate. Qua trnh phan giai xay ra cho en khi chuoi polysaccharide rut
ngan con cach iem phan nhanh 4 n v glucose.
e phosphorylase tiep tuc hoat ong, oligotransferase cat cac n v maltotriose
khoi oan ngan con lai va gan chung vao cac au tan cung khong kh bang lien ket
(1-4). Lien ket (1-6) con lai sau hoat ong cua oligotransferase c cong pha nh -
(1-6)-glucosidase. S phoi hp cua hai enzyme nay lam xuat hien mot mach dai
khong phan nhanh e lai co the chu tac dung cua phosphorylase (hnh IX.2).

Phosphorylase noi bao ton tai hai dang; phosphorylase a co hoat tnh cao va
phosphorylase b khong hoat ong. Qua trnh hoat hoa phosphorylase b thanh
phosphorylase a c thc hien nh hang loat enzyme vi s tham gia cua AMP vong
va nhieu hormone (hnh IX.3).
Hnh IX.2. Phan giai glycogen di tac dung cua phosphorylase (1),
oligotransferase (2) va -(1-6)-glucosidase (3).

Hnh IX.3. S o hoat hoa phosphorylase


Hoa sinh hoc
- 90 -

2.Phan giai cac polysaccharide khac.
Cung nh tinh bot va glycogen, cac polysaccharide khac b phan giai nh nhng
enzyme thuy phan ac hieu.
Cellulose di tac dung cua enzyme cellulase b thuy phan thanh cellobiose.
Enzyme nay co trong nhieu loai nam, mot so con trung va trong da day cua ong vat
an co. Nh he vi sinh vat song cong sinh ay ma nhng ong vat nay tieu hoa c
cellulose. Ngay nay cong nghiep thuy phan cellulose bang cellulase vi sinh vat c
thc hien nhieu nc nham muc ch san xuat ru va nhieu san pham khac, ac
biet la thc an gia suc.
Di tac dung cua -fructosidase inuline b thuy phan thanh -D-fructose, con
rafinose thanh -D-fructose va melibiose, saccharose thanh -D-glucose va -D-
fructose.
Thuy phan cac disaccharide maltose, cellobiose,melibiose va lactose c thc
hien nh cac enzyme ac hieu tng ng -glucosidase, -glucosidase, -
galactosidase, va -galactosidase.
V. CHUYEN HOA TNG HO GIA CAC MONOSE.
S chuyen hoa tng ho gia cac monose c thc hien dang ester phosphate
nh he enzyme cua chu trnh pentosophosphate hoac vi s tham gia cua cac dan xuat
nucleosidediphosphate cua ng (NDPS). Cac dan xuat nay c hnh thanh trong
phan ng gia ester phosphate cua ng vi nucleosidediphosphate nh s xuc tac
cua cac enzyme ac hieu. V du:
Glucoso-1-phosphate + UTP UDP-Glucose + PP
vc
Xuc tac phan ng nay la enzyme uridyldiphosphateglucosopyrophosphorylase.
S chuyen hoa tng ho gia monose dang NDPS c thc hien chu yeu theo
nhng kieu phan ng sau ay:
1.Trao oi (van chuyen) cac nhom glycosyl cua glycosylphosphate:
V du, vi s xuc tac cua hexoso-1-phosphate uridyltransferase say ra phan ng
Gal-1- P + UDP-Glc Glc-1- P + UDP-Gal.
Phan ng nay rat quan trong nham a galactose vao qua trnh trao oi chat.
2.Epimer hoa:
Cac epimerase ac hieu lam bien oi cau hnh cua NDPS. Qua trnh kem theo
cac he thong oxy hoa kh. V du:

Hoa sinh hoc - 91 -

3.Oxy hoa hexose va decarboxyl hoa thanh pentose:
V du: UDP-glucose sau khi b oxy hoa thanh acid UDP-glucuronic b
decarboxyl hoa thanh xylose:

Ben canh s chuyen hoa tng ho gia cac monose, cac dan xuat NDPS con
tham gia trong viec tong hp polysaccharide .
S chuyen hoa tng ho gia cac monose theo chu trnh pentosophosphate se
c xet ti trong muc IV.
VI. GLYCOLYS.
Glycolys la qua trnh phan giai k kh glucose thanh acid lactic. S o tong quat
cua no c trnh bay trong hnh IX.4.
Giai oan th nhat cua glycolis ket thuc sau phan ng E
5
e tao ra cac
triosophosphate. Nhiem vu cua giai oan nay la hoat hoa glucose nh ATP va chuyen
hoa chat nay thanh glyceraldehyde-3-phosphate e tham gia cac phan ng oxy hoa
kh cua giai oan th hai.
Giai oan th hai cua glycolis la giai oan cac phan ng oxy hoa kh va giai
phong ATP. No ket thuc vi s hnh thanh acid lactic (trong cac mo ong vat va mot
so vi khuan) hoac ethanol ( nam men).
Glycolis neu c bat au t glycogen th toan bo qua trnh co ten goi la
glycogenolis. Trong trng hp nay glycogen can c phosphorolis s bo thanh
glucoso-1-phosphate nh glycogenphosphorylase va chat nay se chuyen hoa thanh
glucoso-6-phosphate nh phospho-glucomutase.
Trong so 11 enzyme cua glycolis phosphofructokinase ong vai tro cua enzyme
ieu hoa toan bo qua trnh. Hoat tnh cua no b c che bi ATP va citrate ( nong o
cao) va c kch thch bi ADP va AMP.


Hoa sinh hoc
- 92 -

Hnh IX.4: S o tong quat cua qua trnh glycolys
Hnh .IX.5. C che tac dung cua
glyceraldehyde phosphate
Glyceraldehydephosphate
dehydrogenase cung ong vai tro quan
trong trong viec ieu hoa qua trnh
glycolis. C che tac dung phc tap cua no
c nghien cu kha chi ttiet (hnh IX.5).
Moi mot trong 4 phan di n v giong het
nhau cua enzyme lien ket vi mot phan t
NAD
+
va co mot trung tam hoat ong ma
thanh phan rat quan trong la nhom SH.
Trc tien, enzyme ket hp vi NAD
+
. Sau
o chc aldehyde cua c chat tng tac vi
trung tam hoat ong, tao ra mot
Hoa sinh hoc - 93 -
thiosemiacetal. Giai oan tiep theo la van
chuyen hydro von lien ket vi
glyceraldehyde-3-phosphate en NAD
+

lien ket, kh no thanh NAD.H, ong thi
hnh thanh thioester gia nhom SH cua
enzyme vi nhom carboxyl cua c chat.
Phan t NAD.H van khong tach khoi
enzyme ma lai nhng ien t
va hydro cho phan t NAD
+
t do. Phc he gia c chat va enzyme giai oan nay
c goi la acyl-enzyme. Goc acyl sau o c chuyen t nhom SH cua enzyme en
phan t phosphate vo c e tao ra san pham oxy hoa la acid 1,3-diphosphoglyceric.
Lactate dehydrogenase xuc tac phan ng cuoi cung cua glycolis. Nh a noi
en trc ay, trong c the ong vat bac cao co t nhat 5 dang isoenzyme khac nhau.
Ty le tng oi cua 5 dang nay cung ong gop mot phan quan trong vao viec ieu hoa
glycolis noi rieng va trao oi chat noi chung.
Cac phan ng cua glycolis xay ra trong te bao chat va khong lien quan vi cau
truc te bao. Khi co mat oxy toc o cua no giam sut. Hien tng nay c goi la hieu
ng Pasteur.
Acid lactic san pham cuoi cung cua glycolis trong ieu kien k kh c thai
qua mang te bao ra moi trng nh mot san pham b loai bo. Khi cac te bao c cua
ong vat bac cao hoat ong qua manh trong ieu kien thieu oxy, t c mot lng ln
acid lactic c a vao mau. gan no lai c chuyen hoa thanh glucose. Cam giac
moi met hoac te c la do pH b lech ve pha acid.
Phng trnh tong quat cua glycolis co dang:
Glucose + 2ADP + 2P
vc
2 Lactate + 2ATP + 2H
2
O
Bien thien nang lng t do cua glycolis vi s chuyen hoa glucose thanh 2
lactate co gia tr khong ln (G
o
= -47Kcal). Vi s hnh thanh 2 phan t ATP te bao
tch luy c 2 x 7,3 Kcal = 14,6 Kcal. ieu o co ngha la hieu suat nang lng co
ch cua glycolis bang
14,6 x 100
= 31%
47
sinh vat k kh bat buoc glycolis la nguon nang lng duy nhat. sinh vat k
kh khong bat buoc va sinh vat hieu kh bat buoc glycolis la giai oan au cua s phan
giai glucose. Trong ieu kien co oxy pyruvate khong b kh thanh lactate ma b
decarboxyl hoa oxy hoa thanh acetylcoenzyme A e tiep tuc b phan giai thanh H
2
O
va CO
2
theo chu trnh acid tricarboxylic.

Hoa sinh hoc
- 94 -

C chat cua glycolis ben canh glucose va glucoso-1-phosphate (hnh thanh t
glycogen) con co the la cac monose khac. V du, D-mannose va D-fructose co the
c phosphoryl hoa thanh D-mannoso-6-phosphate va D-fructoso-6-phosphate. D-
fructoso-6-phosphate thc s la san pham trung gian cua glycolis, con D-mannoso-6-
phosphate cung se c chuyen hoa thanh D-fructoso-6-phosphate nh enzyme
phosphomannoso-isomerase. Trong khi o, D-galactose c phosphoryl hoa thanh D-
galactoso-1-phosphate nh galactokinase. Sau o chat nay c epimer hoa thanh
glucoso-1-phosphate vi s tham gia cua UTP nh a trnh bay trong muc V.
Cac pentose c loi cuon vao qua trnh glycolis sau khi chuyen hoa s bo thanh
glyceraldehyde-3-phosphate nh cac phan ng cua chu trnh pentosophosphate.
C chat cua glycolis con co the la glycerine va glycerol-3-phophate von xuat
hien trong qua trnh thuy phan mot so lipid. Glycerine c phosphoryl hoa thanh
glycerol-3-phophate nh glycerokinase. Sau o glycerophosphate dehydrogenase oxy
hoa chat nay thanh dioxyacetonephosphate, mot trong nhng san pham trung gian cua
glycolis.
Qua trnh len men ru xay ra trong te bao nam men vi s tham gia cua 10
enzyme au cua glycolis va 2 enzyme bo sung la pyruvate decarboxylase va
alcoholdehydro-genase.
Enzyme pyruvate decarboxylase xuc tac cho phan ng khong thuan nghch
Pyruvate Acetaldehyde + CO
2

C che tac dung cua enzyme nay c trnh bay trong hnh IX.6. Trc tiep tham
gia trong hoat ong xuc tac cua enzyme la coenzyme thiaminepyrophosphate (dan
xuat cua vitamine B
1
). Hoat ong cua no can co Mg
2+
.
Qua trnh decarboxyl
hoa acid pyruvic c thc
hien qua mot loat giai oan.
Trc tien, nguyen t carbon
cua pyruvate ket hp vi C
2

cua vong thiazol cua TPP
trang thai lien ket vi enzyme,
lam xuat hien dan xuat 2--
lactyl cua coenzyme. Dan xuat
nay sau o b decarboxyl hoa
thanh dan xuat 2-oxyethyl
(acetaldehyde hoat ong).
Cuoi cung, nhom oxyethyl
tach khoi coenzyme dang
acetaldehyde t do. Chat nay
Hoa sinh hoc - 95 -

sau o se b kh thanh ethanol
nh alcoholdehydro-genase:
Acetaldehyde + NADH +
H
+
Ethanol + NAD
+

Phng trrnh tong quat
cua len men ru v vay se la:
Glucose + 2ADP + 2P
vc

2 Ethanol + 2CO
2
+
2ATP.
VII. CHU TRNH PENTOSOPHOSPHATE
Chu trnh pentosophosphate (con c goi la con ng phosphogluconate) la
mot trat t phan ng xay ra trong te bao chat song song vi glycolis, bao gom 2 phan
ng oxi hoa kh va 9 phan ng chuyen hoa tng ho gia cac ester phosphate cua
triose, tetrose, pentose, hexose va heptose (hnh IX.7).
Dehydrogen hoa va decarboxyl hoa 6 phan t glucoso-6-phosphate tao ra 6
phan t pentoso-5-phosphate. Nhng phan t pentosophosphate nay trai qua cac phan
ng chuyen hoa tng ho do cac enzyme trancetolase, transaldolase aldolase,
isomerase va epimerase xuc tac. San pham trung gian la cac ester phosphate cua
monose cha t 3 en 7 nguyen t carbon. Nhng ester nay tao ra 4 phan t fructoso-
6-phosphate va 2 phan t glyceralde-hyde-3-phosphate. Hai phan t nay se ngng tu
e tao ra phan t fructoso-6-phosphate th 5 hoac tiep tuc b phan giai theo cac phan
ng glycolis. Cac phan t fructoso-6-phosphate se c ong phan hoa thanh glucoso-
6-phosphate va gia nhap vao von glucoso-6-phosphate chung cua te bao. Von nay co
the c s dung cho cac phan ng glycolis hoac cac phan ng cua chu trnh
pentosophosphate.

Hnh IX.6. C che tac dung cua pyruvate
decarboxylase
Hoa sinh hoc - 96 -

Phng trrnh tong quat cua chu trnh pentosophosphate la:
Hnh IX.7.Chu trnh pentosophosphate
1. Glucoso-6-phosphatye dehydrogenase; 2. 6-phosphogluconate
dehydrogenase; 3.Phosphopentoepimerase; 4.Phoaphopentoisomerase;
5,7.Transcetolase; 6. Transaldollase; 8. Triosophosphate isomerase; 9.
Aldolase; 10. Fructodiphosphatase; 11. Phosphoglucoisomerase.
6 Glucoso-6-phosphate + 12 NADP
+
5 Glucoso-6-phosphate + 6CO
2

+
12 NADP.H + 12 H
+
+ P
vc

Hoac gon hn:
Glucoso-6-phosphate + 12 NADP
+
6CO
2
+ 12 NADP.H + 12 H
+
+ P
vc. .
Tuy nhien, theo con ng nay glucose thng khong b oxy hoa hoan toan
thanh CO
2
. Cac phan ng cua chu trnh nay khong the c s dung lam nguon nang
lng. Do xay ra trong te bao chat nen oxy hoa NADP.H (von hnh thanh trong chu
trnh) khong the tao ra ATP mot cach trc tiep nh nhng phan t NAD.H va FAD.H
2

trong ty the. Chc nang chu yeu cua chu trnh pentosophosphate la tao ra NADP.H e
cung cap cho cac phan ng kh cua cac qua trnh sinh tong hp. No con la nguon cung
cap riboso-5-phosphate e tong hp nucleotide, acid nucleic va nguon erytroso-4-
phosphate e tong hp cac aminoacid vong thm trong cac c the t dng.
Neu khong c dung cho cac muc ch sinh tong hp, cac chat trao oi trung
gian cua chu trnh se c chuyen hoa thanh glyceraldehyde-3-phosphate va
fructoso-6-phosphate e tham gia qua trnh glycolis.
Theo hng ngc lai, mot so phan ng cua chu trnh pentosophosphate c
dung e co nh CO
2
trong quang hp.
VIII. OXY HOA HIEU KH GLUCID.
1.Decarboxyl hoa oxy hoa acid pyruvic.
sinh vat hieu kh qua trnh d hoa glucid thng c chia lam hai giai oan:
giai oan k kh (glycolis) vi s hnh thanh acid pyruvic va giai oan hieu kh, trong
o acid pyruvic se b oxy hoa hoan toan thanh CO
2
va H
2
O. Phan ng au tien cua
giai oan sau la decarboxyl hoa oxy hoa acid pyruvic thanh acetylcoenzyme A vi s
xuc tac cua pyruvate dehydrogenase:
CH
3
-CO-COOH + NAD
+
+ CoA-SH CH
3
-CO-SCoA + CO
2
+ NAD.H + H
+

Qua trnh phan ng xay ra nh mo ta trong hnh IX.8.

Hoa sinh hoc
- 97 -

Pyruvatedehydrogenase la mot he thong enzyme phc tap gom 3 enzyme va 5
coenzyme coenzyme tap hp lai thanh mot c cau thong nhat, trong o moi enzyme
va coenzyme thc hien mot chc nang xac nh. Hoat tnh cua cua he thong enzyme
nay b c che khi ham lng ATP trong te bao vt qua gii han nhat nh. Co ngha
la Pyruvatede-hydrogenase la mot enzyme ieu hoa. No ong vai tro quan trong trong
viec chi phoi hoat ong cua chu trnh acid tricarboxilic va cua cac qua trnh lien quan
khac.

Hnh IX.8. S o cac giai oan cua phan ng decarboxyl hoa oxy hoa acid
pyruvic.
E1: Pyruvate decarboxylase co coenzyme la thyamine pyrophosphate (TPP). E2-
dihydrolipoyltransacetylase co coenzyme la acuid lipoic;
E3:Dihydrolipoyldehydrogenase co coenzyme la FAD.
2. Chu trnh acid tricarboxylic (Chu trnh Krebs)
Acetylcoenzyme A tiep tuc b oxy hoa thanh kh carbonic theo mot c che co ten
la chu trnh acid tricarboxylic hay chu trnh Krebs (hnh IX.9).

Hoa sinh hoc - 98 -

Hnh IX.9. Chu trnh acid tricarboxylic. 1:Citrate syntase; 2.3: Aconitase;
4: Isocitrate dehydrogenase; 5: Cetoglutarate dehydrogenase; 5a:Succinate
thiokinase; 6: Succinate dehydrogenase; 7: Fumarase; 8: Malate dehydrogenase.

Citrate syntase, isocitrate dehydrogenase va fumarase la nhng enzyme mang
tnh chat cua enzyme ieu hoa. Hoat tnh cua chung gop phan chi phoi toc o cua chu
trnh. Hoat tnh cua nhng enzyme nay b c che bi ATP va NAD.H. Tuy nhien, ve
phng dien nay isocitrate dehydrogenase co vai tro quan trong hn ca. Khi ham
lng ADP trong te bao tang len, toc o oxy hoa isocitrate t ong tang, lam tang toc
o cua toan bo chu trnh. Ngc lai, khi tang toc o cua chu trnh, ham lng ATP
trong te bao tang len, lam cho hoat ong cua isocitrate dehydrogenase b c che. S
tch luy NAD.H trong ty the cung co tac dung tng t.
ang lu y hn ca ve phng dien tch luy nang lng la phan ng -
cetoglutarate dehydrogenase. Phc he enzyme nay rat giong vi he thong pyruvate
dehydrogenase. Hoat ong cua no cung oi hoi cac coenzyme TPP, acid folic, CoA-
SH, NAD
+
va FAD. Vi s tham gia cua nhng coenzyme nay -cetoglutarate b oxy
hoa thanh suxinyl-coenzyme A. Suxinyl-coenzyme A la mot tioester cao nang. giai
oan ke tiep nang lng tch luy trong no c dung e tong hp GTP t GDP va P
vc

nh s xuc tac cua enzyme suxinyl thiokinase ( suxinyl-CoA-synthetase). GTP sau o
nhng goc phosphate tan cung cho ADP e tao ra phan t ATP.
Kieu tong hp ATP nay thng c goi la phosphoryl hoa oxy hoa mc o c
chat e phan biet vi cac phan ng phosphoryl hoa oxy hoa trong chuoi ho hap (muc
IX).
Toan bo cac enzyme cua chu trnh acid tricarboxylic cung nh cac enzyme cua
chuoi van chuyen ien t en oxy phan t eu nh v trong ty the cau truc c
menh danh la Tram nang lng cua te bao. Mot so enzyme cua chu trnh nay cung
co mat trong phan oan hoa tan cua te bao chat, tuy nhien chung tac dung len cac con
ng trao oi chat khac.
3. Y ngha cua chu trnh acid tricarboxylic.
Qua hnh IV.9 ta thay rang mot vong cua chu trnh dung nap mot phan t acetyl-
CoA va oxy hoa chat nay thanh hai phan t CO
2
, ong thi tao ra mot phan t ATP va
4 coenzyme dang kh (3 NAD.H va 1 FAD.H
2
). Nhng phan t coenzyme nay se
tao ra so nang lng ang ke khi b oxy hoa trong chuoi ho hap.

Hoa sinh hoc - 99 -
Nh vay, cac phan ng cua chu trnh acid tricarboxilic la nguon cung cap nang
lng quan trong cua te bao. Tuy nhien no con lien quan ca vi cac qua trnh ong
hoa vi t cach la nguon cung cap nguyen lieu va nang lng cho hang loat cac phan
ng sinh tong hp. Mot so san pham trung gian cua chu trnh nh nhng phan ng lien
he c dung e tong hp glucose, acid beo aminoacid va nhieu hp chat khac. Phan
ln nhng phan ng nay co tnh thuan nghch nen chung cung c s dung e tao ra
cac san pham trung gian cua chu trnh t nhng hp chat khong phai glucid. Do ac
iem nay chu trnh acid tricarboxilic c xem la trung tam cua trao oi chat te bao.
4. Cac phan ng bu ap.
Thong thng, nong o cua cac san pham trung gian cua chu trnh acid
tricarboxilic trong ty the c gi mc rat on nh. Trong viec gi the on nh nay
ngoai vai tro cua cac enzyme ieu hoa nh pyruvate dehydrogenase isocitrase
dehydrogenase v.v... con co s tham gia cua nhng phan ng enzyme ac biet chuyen
lam nhiem vu bo sung nguon d tr cac san pham trung gian cua chu trnh ma ngi ta
goi la nhng phan ng bu ap. Trong so cac phan ng loai nay quan trong nhat la phan
ng carboxyl hoa acid pyruvic thanh acid oxaloacetic do enzyme pyruvate
carboxylase xuc tac:
Mg
2+
Pyravate + CO
2
+ ATP Oxaloacetate + ADP + P
vc

Khi chu trnh hoat ong trong ieu kien thieu oxaloacetate hoac thieu cac san
pham trung gian khac, phan ng nay c kch thch lam cho nguon d tr
oxaloacetae tang len. Pyruvate carboxylase la mot enzyme ieu hoa. Toc o cua
phan ng thuan tang len ang ke khi trong te bao co d acetyl-CoA; ngc lai, khi
thieu acetyl-CoA, toc o cua phan ng rat thap. Trong gan va than cua ong vat bac
cao ay la phan ng bu ap quan trong nhat.
nhng mo khac chc nang bu ap c thc hien bi malico-enzyme (malate
dehydrogenase) va phosphoenolpyruvate carboxylase xuc tac cac phan ng tng ng
sau ay:
L.Malate + NADP
+
Pyruvate + CO
2
+ NADP.H + H
+
va
Phosphoenolpyruvate + CO
2
+

IDP Oxaloacetate + ITP
thc vat va vi sinh vat chc nang bu ap con c thc hien bi chu trnh
glyoxylate (chng 5).
IX. PHOSPHORYL HOA OXY HOA.
Trong trat t cac phan ng d hoa glucose ma ta a xem xet tren ay co 4 phan
t ATP c hnh thanh trong cac phan ng lien hp vi cac phan ng oxy hoa
aldehyde 3-phosphoglyceric va acid -cetoglutaric. So phan t ATP nay mi ch la
mot phan nho nang lng san sinh trong toan bo qua trnh phan huy mot phan t

Hoa sinh hoc - 100 -
glucose thanh kh carbonic va nc. So nang lng con lai mot phan c giai phong
dang nhiet von can e thuc ay qua trnh phan ng, mot phan khac c tch luy
trong cac coenzyme kh NAD.H va FAD.H
2
von co the kh tieu chuan rat cao va do
o la nhng hp chat rat giau nang lng . Nh ta a thay, co tat ca 12 phan t thuoc
loai nay c hnh thanh khi phan huy mot phan t glucose, bao gom 2 NAD.H trong
glycolis, 2 NAD.H trong phan ng decarboxyl hoa oxy hoa acid pyruvic thanh acetyl-
CoA va 6 NAD.H cung 2 FAD.H
2
trong chu trrnh acid tricarboxylic.
Trong ieu kien hieu kh nhng hp chat giau nang lng nay se chuyen ien t
cua mnh cho oxy khong kh e khoi phuc lai trang thai oxy hoa NAD
+
va FAD. Tuy
nhien, oxy khong trc tiep nhan ien t t cac coenzyme kh. Cac ien t cung vi
proton von gan lien vi chung c a en chat nhan cuoi cung qua mot chuoi chat
van chuyen trung gian nh v tren mang trong cua ty the, co ten goi la chuoi ho hap.
Nang lng giai phong trong qua trnh van chuyen ien t nay sau o c he enzyme
ATP-ase cung nh v tren mang trong cua ty the dung e tong hp ATP trong qua
trnh phosphoryl hoa oxy hoa.
S hieu biet cua con ngi hien nay ve phosphoryl hoa oxy hoa (va ca
phosphoryl hoa quang hp) da tren gia thuyet hoa tham thau do Peter Mitchell e
xuat nam 1961, theo o hieu so nong o proton gia hai pha cua mang trong cua ty
the (va mang thylacoid cua luc lap) la ong lc quan trong cho qua trnh sinh tong hp
ATP t ADP va P
vc
.
Co 3 iem giong nhau c ban gia phosphoryl hoa oxy hoa va phosphoryl hoa
quang hp:
1/ Ca hai qua trnh eu gan vi dong ien t xuyen qua mot chuoi cac chat van
chuyen trung gian dang kh gan vi mang, bao gom cac hp chat quinone,
cytochrome va cac protein cha sat lu huynh;
2/ Nang lng t do giai phong do dong ien t i xuong gan lien vi s van
chuyen proton i len xuyen qua he thong mang khong tham oi vi proton. S phoi
hp nay giup tch luy mot phan nang lng dang the ien hoa xuyen mang;
3/ Dong proton xuyen mang theo chieu giam nong o cua chung xay ra xuyen
qua cac kenh ac hieu, dan en s cung cap nang lng cho sinh tong hp ATP.
Trong muc nay chung ta xem xet trc qua trnh phosphoryl hoa oxy hoa, bat
au bang s mo ta cac thanh phan cua chuoi van chuyen ien t tren mang trong cua
ty the, trat t cac chat van chuyen c to chc thanh cac phc he chc nang e hoat
ong, ong thi, se xet ti c che hoa tham thau ma nh o s van chuyen ien t
trong chuoi ho hap dan en sinh tong hp ATP.
Ty the c phan cach cac c quan t khac cua te bao bi lp mang kep. Mang
ngoai cho phep cac phan t nho va cac ion xuyen qua. Cac kenh xuyen mang cau tao
t protein cho phep phan ln cac phan t vi M < 5000 qua lai de dang. Trong khi o
mang trong khong xuyen tham oi vi phan ln cac phan t nho va ion, ke ca ion H
+
.

Tren mang nay nh v cac thanh phan cua chuoi ho hap va phc he enzyme lam
nhiem vu tong hp ATP.
Trong thanh phan cua chuoi ho hap ngoai NAD va cac flavoprotein con co 3
nhom van chuyen ien t khac. o la benzoquione k nc (ubiquinone) va hai nhom
protein cha sat la cac cytochrome va Fe-S-protein.
Trong tat ca cac phan ng xay ra tren chuoi ho hap cac ien t di chuyen t
NAD.H, succinate hoac mot so chat cho ien t s cap khac, xuyen qua cac
flavoprotein, ubiquinone Fe-S-protein va cac cytochrome e cuoi cung en vi O
2

(hnh IX.10).
Da vao the kh cua chung co the sap xep cac chat van chuyen theo trat t:
NAD.H, UQ, cytochrome b, cytochrome c
1
, cytochrome c, cytochrome a+a
3
.
X ly
nhe mang
trong cua ty
the bang chat
tay ra, cho
phep phat
hien 4 phc he
van chuyen
ien t, moi
phc he co
thanh phan rat
ac trng (
bang IX.1).
Moi phc he
co kha nang
xuc tac van
chuyen ien t
qua mot khu vc cua chuoi ho hap.
Cac phc he I va II xuc tac van chuyen ien t en ubiquinone t hai chat cho
khac nhau la NAD.H (phc he I) va succinate (phc he II). Phc he III van chuyen
ien t t ubiquinone en cytoxhrome c, con phc he IV tiep tuc van chuyen ien t
t cytochrome c en O
2
.
Bang IX.1. Thanh phan protein cua chuoi van chuyen ien t trong ty the
Phc he enzyme
Khoi lng
(kDa)
So lng phan di
n v
Cac nhom them
I. NAD.H-
dehydrogenase
850 > 25 FMN,Fe-S
II. Succinate
dehydrogenase
140 4 FAD, Fe-S
Hnh IX.10. S o mo ta chuoi ho hap va hoat ong cua qua
trnh phosphoryl hoa oxy hoa.

III. Ubiquinone,
cytochrome c-
oxydoreductase
Cytochrome c

250

13

10

1

Heme, Fe-S

Heme
IV. Cytochrome c
oxydase
160 6-13 Heme, Cu
A
,Cu
B
Ngoai 4 phc he tren, tren mang trong cua ty the con co he enzyme sinh tong
hp ATP, ky hieu la phc he F
o
F
1
, hay oi khi con c goi la phc he V.
Qua hnh IX.10 ta co the thay ro cac ien t at en UQ qua cac phc he I va II.
UQ.H
2
lam nhiem vu nh mot chat van chuyen linh ong oi vi ien t va proton. No
chuyen ien t cho phc he III e sau o phc he III chuyen tiep cho cytochrome c.
Phc he IV chuyen ien t t cytochrome c en O
2
. Dong ien t xuyen qua cac phc
he I, III va IV kem theo dong proton t c chat cua ty the i ra khoang khong gian
gia 2 mang tao ra gradient proton xuyen mang ong lc e he enzyme ATP-ase s
dung trong viec sinh tong hp ATP sau o.
Nh vay, nh hoat ong cua chuoi ho hap ma toan bo nang lng tch luy trong
cac coenzyme kh NAD.H va FAD.H
2
a c giai phong. Phan ln so nang lng o,
nh ta a thay, a chuyen hoa thanh nang lng cua gradient proton gia hai pha cua
mang trong cua ty the. Gradient nay hnh thanh c la nh tnh khong xuyen tham
cua ion H
+
qua mang nay. Ngan trong b mat H
+
se tnh ien am, con ngan ngoai co
nhieu ion H
+
nen tch ien dng. Ket qua xuat hien mot hieu so ien the. Phc he
F
o
F
1
la ni duy nhat cho phep ion H
+
quay ngc t ngan ngoai vao ngan trong. Khi
dong ien ( ay la dong H
+
ch khong phai la dong e
-
) chay qua, phc he nay se
c cap nang lng e tong hp ATP t ADP va P
vc
. Nh vay, nh co s hoa hp
gia phc he F
o
F
1
vi chuoi ho hap ma nang lng giai phong trong cac phan ng oxy
hoa NAD.H va FAD.H
2
co the c dung e tong hp ATP. V the kieu tong hp ATP
nay c goi la phosphoryl hoa oxy hoa trong chuoi ho hap.
Ngoai tac dung lam nguon nang lng trc tiep e tong hp ATP, gradient
proton con c s dung e bm cac phan t ATP t trong c chat cua ty the ra te bao
chat.
Tren c s mc chenh lech nang lng gia NAD.H, FAD.H
2
vi dang oxy hoa
tng ng (NAD
+
va FAD) ngi ta cho rang khi nhng iien t cua mnh cho oxy
phan t qua chuoi ho hap cho phep tao ra 3 ATP t NAD.H va 2 ATP t FAD.H
2
.
Rieng cac phan t NAD.H hnh thanh trong qua trnh glycolis von xay ra trong te bao
chat ch cho phep tao ra so phan t ATP tng ng vi FAD.H
2
trong ty the. o la
do ket qua cua mot c che van chuyen cac ng lng kh t te bao chat vao ty the
co ten la c che con thoi glycerophoosphate. Theo c che nay NAD.H cua te bao chat
c dung e kh dioxyacetone phosphate thanh glycerophosphate. Sau o
glycerophosphate i vao ty the. Tai o vi s tham gia cua FAD no b oxy hoa ngc
lai thanh dioxyacetone phosphate, con FAD b kh thanh FAD.H
2
. Ca hai phan ng

eu c xuc tac bi glycerophosphate dehydrogenase nhng vi s tham gia cua hai
coenzyme khac nhau. Ket qua la cac ng lng kh trong te bao chat dang
NAD.H c chuyen vao ty the dang FAD.H
2
.
Nh hoat ong cua chuoi ho hap lien hp vi phc he F
o
F
1
ma nang lng t do
giai phong t mot phan t glucose trong qua trnh glycolis co the dan en s hnh
thanh them 4 phan t ATP, nang tong so ATP cua giai oan nay len 6 phan t. Trong
khi o giai oan hieu kh tiep theo ngoai 2 ATP xuat hien trc tiep trong chu trnh
acid tricarboxilic con hnh thanh them (6x3) + (2x2) = 28 phan t ATP na. Do o khi
oxy hoa hoan toan mot phan t glucose thanh kh carbonic va nc tao ra c 36
phan t ATP, trong o 6 phan t giai oan k kh va 30 phan t giai oan hieu kh.
Trong moi phan t ATP tch luy 7,3 Kcal. Nh vay trong 36 phan t ATP tch luy c
7,3 x 36 = 263 Kcal.
Trong khi o bien thien nang lng t do cua phan ng C
6
H
12
O
6
+ 6O
2

6CO
2
+ 6H
2
O la 686 Kcal. T o ta co the thay ro hieu suat tch luy nang lng cua
phan ng nay la 263 x 100 / 686 = 38%.
X. QUANG HP
1. Phng trnh tong quat cua quang hp.
Moi c the song ch ton tai khi c lien tuc cung cap nang lng t ben ngoai. Dang
nang lng duy nhat ma c the song co the s dung c e thc hien cong la nang lng ho a
hoc. Nguon nang lng nay c khai thac t hai hng: nang lng hoa hoc cua cac hp chat
vo c (hng trai at) va nang lng anh sang (hng vu tru). Quang hp chnh la qua trnh
bien oi nang lng anh sang thanh nang lng hoa hoc di dang cac hp chat hu c, c
thc hien bi cac c the va te bao co cha chat diep luc.
Theo cac nh luat cua nhiet ong hoc, moi phan ng t phat eu kem theo giam
nang lng t do va tang entropy cua he thong, co ngha la lam cho he thong tr nen
mat trat t. Tnh to chc cua mot he thong ch co the c nang cao khi no c cung
cap them nang lng, tao ieu kien e phan ng co the xay ra ngc vi gradient
nhiet ong hoc. Nh co quang hp ma nang lng anh sang c chuyen hoa thanh
nang lng hoa hoc dang de s dung e cung cap cho cac phan ng can s dung
nang lng nham tao lap, duy tr va nang cao tnh to chc cua te bao va c the.
Sau khi nang lng anh sang a c cac phan t diep luc (chlorophyll) hap thu
va bien oi thanh nang lng hoa hoc, no se c dung e kh CO
2
thanh cac hp
chat hu c, ma chu yeu la glucose. V vay, phng trnh tong quat cua quang hp co
the viet di dang:
h
6CO
2
+ 6H
2
O C
6
H
12
O
6
+ 6O
2


Tuy nhien, khong phai qua trnh quang hp nao cung giai phong oxy. Mot so vi
khuan quang hp khong dung nc ma dung mot so hp chat khac e lam chat cho ien t
trong qua trnh quang hp. V vay quang hp nhng c the nay khong giai phong oxy. V du:
h
2CH
3
-CHOH-CH
3
+ CO
2
[CH
2
O] + CH
3
-CO-CH
3

h
suxinat + CO
2
[CH
2
O] + Fumarate;
h
2 H
2
S + CO
2
[CH
2
O] + 2S.
Thong qua cac phng trnh tren, co the viet phng trnh quang hp dang khai quat
hn nh sau:
h
2H
2
D + CO
2
[CH
2
O] + 2D
trong o DH
2
la chat cho ien t, con D la dang oxy-hoa cua no.
Mat khac, trong cac loai te bao quang hp khac nhau, tuy chat nhan ien t (A)
chu yeu la CO
2
, nhng trong mot so trng hp A co the la cac hp chat khac, v du
thc vat bac cao A co the la nitrate, vi sinh vat A co the la N
2
. Tham ch mot so c
the quang hp co the s dung ca H
+
lam chat nhan ien t. Trong trng hp nay H
+

se b kh thanh H
2
.
Nh vay, co the viet phan t quang hp dang chung nhat nh sau:
h
DH
2
+ A AH
2
+ D

Du qua trnh quang hp xay ra vi s tham gia cua bat ky chat cho va chat
nhan ien t nao, dong ien t t chat cho en chat nhan hnh thanh di tac dung cua
anh sang mat tri bao gi cung co hng ngc lai vi hng bnh thng cua gradient
the kh tieu chuan ac trng cho he thong chat nhan - chat cho o, tc hng ve pha
co gia tr am cua the kh tieu chuan cao hn. Bang cach o nang lng anh sang c
tch luy lai trong phan t chat nhan dang the oxy-hoa kh.
Cac nghien cu chi tiet qua trnh quang hp cho thay rang e tong hp mot
phan t glucose can s dung 12 phan t nc, va trong so san pham, can phai ke en 6
phan t nc khac vi 12 phan t nc a s dung. V vay, phng trnh tong quat ay
u cua qua trnh quang hp thc vat bac cao can phai viet di dang:
h
6CO
2
+ 12H
2
O 6O
2
+ C
6
H
12
O
6
+ 6H
2
O

Bien thien nang lng t do tieu chuan (G
o
') cua phan ng tren ay bang -
1300 KCal/mol. V nang lng tch luy trong phan t glucose la 686 KCal/mol, nen e
tch luy c so nang lng nay te bao quang hp can phai thu nhan khoang
1800KCal/mol nang lng anh sang.
2. Khai niem ve tch chat hai giai oan cua quang hp.
Tren c s thc nghiem ngi ta a xac nh c rang qua trnh quang hp bao
gom hai loai phan ng: phan ng sang lien quan trc tiep vi viec s dung nang lng

anh sang, va phan ng toi, trong o CO
2
b kh thanh cac hp chat hu c. Khai niem
ve tnh hai pha cua quang hp c uc rut t ket qua th nghiem cua Hill (1937).
Bang cach chieu sang che pham phi te bao cha chlorophyll thu c t te bao quang
hp trong ieu kien ton tai chat nhan ien t nhan tao, v du kali ferricyanate (A),
oxy phan t se c giai phong, ong thi chat nhan ien t b kh theo phng trnh:
h
2H
2
O + 2A 2AH
2
+ O
2

ieu ac biet quan trong la phan ng khong oi hoi s tham gia cua CO
2
. Nh
vay, Hill a cho thay kha nang tach qua trnh giai phong O
2
ra khoi qua trnh kh CO
2

.
Phan ng tren ay ngay nay c goi la phan ng Hill, con chat A c goi la
thuoc th Hill. Ngi ta cung a phat hien c rang thuoc th Hill trong te bao la
NADP
+
. Trong ieu kien luc lap c chieu sang, no se nhan ien t e b kh thanh
NADP.H, ong thi giai phong oxy phan t. Nh vay, phan ng Hill trong te bao co
dang:
h
2H
2
O + 2NADP
+
2NADP.H + 2H
+
+ O
2

Tren c s cua phan ng nay ngi ta cho rang mot trong nhng san pham cuoi
cung cua pha sang la mot tac nhan kh nao o (co the la NADP.H) ma sau o trong
pha toi se c s dung e kh CO
2
.
Khai niem ve tnh chat hai pha cua quang hp c tiep tuc cung co nh Arnon
phat hien qua trnh phosphoryl-hoa quang hp nam1954. Ong nhan thay rang khi
chieu sang luc lap trong ieu kien co mat ADP va phosphate vo c th ATP se c
tong hp ma cung khong can co s tham gia cua CO
2
.
Tren c s cua cac th nghiem noi tren ngi ta i en ket luan rang giai oan
au cua qua trnh quang hp nang lng anh sang khong nhng c dung e kh
NADP
+
ma con co tac dung thc hien phan ng phosphoryl-hoa ADP thanh ATP.
NADP.H va ATP hnh thanh pha sang se c s dung trong pha toi e kh CO
2
va
cac chat nhan ien t khac.
3. Vai tro cua nang lng anh sang oi vi quang hp.
Pha sang cua quang hp bao gom hai qua trnh lien quan mat thiet vi nhau: van
chuyen ien t va phosphoryl-hoa. Moi quan he nay tng t nh moi quan he gia
cac qua trnh xay ra tren mang ty the. S khac biet ay la cho dong ien t chuyen
ngc vi gradient the kh tieu chuan. S d co dong ien t ngc nay la nh kha
nang cua chlorophyll hap thu nang lng anh sang.
a/ Anh sang va chlorophyll.
Anh sang ch bao gom mot phan nho cua quang pho bc xa ien t (hnh IX.11).
Moi loai bc xa c ac trng bi bc song va sc cha nang lng. Hai ac trng
nay co tng quan nghch vi nhau: bc song cang dai sc cha nang lng cang
nho. Trong pham vi mot dai hep ma mat ngi nhn thay c cac song anh sang ngan
nhat gay cam giac mau tm va dai nhat - mau o.

Phan ln nang lng mat tri c bc xa dang photon (lng t). Nang lng
cua photon c tnh bang cong thc:
C 286.000
E = Nh = Nh = KCal

trong o E la nang lng tnh bang KCal cua mot "mol photon", hay mot
enstein; N la so Avogadro (bang 6,023.10
23
); la tan so (so dao ong cua song ien
t trong 1 giay); h la hang so Plank (bang 1,58 x 10
-34
Calo/s; C la toc o anh sang
(bang 3.10
10
cm/s); la bc song anh sang o bang .

T cong thc tren co the tnh c nang lng trung bnh cua cac tia sang nhn
thay nh sau: tia o: 40,86KCal/enstein; tia vang: 47,67 KCal/enstein; tia xanh: 57,20
KCal/enstein va tia tm: 71,50 KCal/enstein.
Hnh IX.11 Cac bo phan cua quang pho ien t

HNH IX.12. QUANG PHO HAP THU (A)
VA QUANG PHO HOAT ONG (B) CUA
Kha nang cua
te bao quang hp s
dung anh sang vi loai
bc song nao la phu
thuoc vao s co mat
cua cac sac to quang
hp. Trong so cac sac
to quang hp nay quan
trong nhat la
chlorophyll. Bang cach
xac nh quang pho
hap thu co the biet moi
loai sac to quang hp
hap thu nhieu nhat
nhng tia nao. Hnh
IX.12A gii thieu
quang pho hap thu cua
chlorophyll

a, chlorophyll b va carote-noid. Ta co the thay ro chlorophyll a hap thu chu yeu
cac tia vi cac bc song tm, lam va o. Cac tia
luc, vang va da cam ch c chlorophyll a hap thu rat t. Mot so vung cua quang pho
mac du c chlorophyll hap thu rat t, nhng van co hieu lc kha cao oi vi quang
hp, lam cho pho hoat ong cua quang hp c ni rong (hnh IX.12B). o la do cac
tia tai nhng vung quang pho nay c cac sac to khac hap thu (carotenoid va cac
dang chlorophyll khac) va chuyen nang lng cho chlorophyll a. Nhng sac to phu tr
nay a lam cho thc vat s dung anh sang vi bc song a dang hn la nhng tia ch
c chlorophyll hap thu.
Trong luc lap cua te bao quang hp cay xanh chlorophyll va cac sac to quang hp phu
tr c sap xep thanh nhom chc nang goi la cac n v quang hp. Moi n v quang h p
cha khoang 300 phan t sac to bao gom chlorophyll a, chlorophyll b, carotenoid v.v... Trong
so cac phan t sac to nay cua moi n v quang hp co mot phan t sac to khac biet vi tat ca
cac phan t con lai. o la mot dang ac biet cua chlorophyll a hoat ong nh mot trung tam
phan ng. Cac phan t sac to khac hoat ong nh nhng anten thu nhan nang lng anh sang.

HNH IX.13. S O MO TA CAC MC O KCH
ONG IEN T KHAC NHAU TRONG PHAN T

Khi mot photon tac ong len mot phan t chlorophyll va c hap thu, nang
lng cua no c chuyen cho mot ien t cua phan t sac to, lam cho ien t b kch
ong nay chuyen len mc nang lng cao hn va tng oi khong on nh (hnh
IV.13). Khong phai photon nao cung lam c viec nay, bi v cac ien t co nhng
mc nang lng xac nh. Photon can phai co u nang lng mi co the kch ong
ien t cua sac to t trang thai c ban len mot mc nang lng cao hn nao o. Cac
photon nay con c goi la exiton . Khong phai moi ien t eu oi hoi mot gia tr
nang lng nh nhau e c nang len mc kch ong. V du, mot ien t t mc o
c ban thap hn (hnh IX.13B) cung co the b kch ong len cung mc nang lng M
(nh trong hnh IX.13A) nh mot photon vi nang lng tng ng e sau o tham
gia vao qua trnh quang hp. Photon giau nang lng hn va bc song ngan hn nay

ai dien cho vung tm-lam cua quang pho hap thu, trong khi photon ngheo nang lng
hn trong trng hp A ai dien cho vung o.
ien t b kch ong, va do o khong ben, se t quay ve trang thai c ban cua no
hau nh ngay tc khac va giai phong nang lng hap thu c. Chlorophyll chiet xuat
trong ong nghiem nhanh chong e mat nang lng ma no chiem gi c bang cach
phat lai dang anh sang huynh quang. o la v cac phan t chlorophyll tach biet khoi
n v quang hp khong con kha nang bien hoa nang lng anh sang thanh nang lng
hoa hoc. Nhng trong luc lap, khi ma nang lng anh sang a nang mot ien t trong
mot phan t anten len mc nang lng cao hn, trang thai kch ong se c chuyen
t phan t sac to nay en phan t sac to khac va cuoi cung at en phan t sac to trung
tam (bay exiton) e b phan t sac to nay chiem oat. Trang thai kch ong b chiem
oat tai ay la nh nang lng can e kch ong phan t sac to trung tam thap hn so
vi nang lng can e kch ong cac phan t sac to anten.
Nh vay, moi khi trang thai kch ong at ti trung tam phan ng, no khong de
g tron thoat c. Trong phan t sac to trung tam trang thai kch ong ien t giau
nang lng khong quay ve trang thai c ban nh trong cac sac to khac, thay vao o no
c chuyen cho mot phan t chat nhan va i qua mot loat cac phan ng enzyme e
tch luy nang lng nay trong NADP.H va ATP.
b/ Khai niem ve hai he thong quang ho a va hai kieu phan ng sang.
Nhieu dan lieu thc nghiem cho thay trong qua trnh quang hp thc vat bac
cao ton tai hai loai phan ng sang.
Trong luc lap cua thc vat co hai loai chlorophyll khac nhau, trong khi o vi
khuan quang hp, ma qua trnh quang hp xay ra khong kem theo giai phong oxy, th
ch co mot loai chlorophyll. Hn the na, cac th nghiem cua Emerson cho thay rang
hieu suat quang hp cua thc vat trong vung anh sang o ke t bc song 680nm tr
len giam sut nhanh chong. Tuy nhien, neu ong thi vi cac tia 680nm chieu te bao
bang cac tia co bc song ngan hn ( = 650nm) th hieu suat quang hp oi vi tia
680nm lai c khoi phuc en gia tr bnh thng. ac biet la cng o quang hp khi
chieu ong thi hai loai tia 680 va 650nm se co gia tr ln hn so vi tong gia tr
cng o quang hp khi hai loai tia nay c chieu rieng biet. Tren c s o ngi ta
cho rang e at hieu suat quang hoa cao nhat, can phai xay ra ong thi hai loai phan
ng sang khac nhau, ma c s cua hai loai phan ng nay la hai he thong quang hoa
khac nhau ton tai trong luc lap cua thc vat bac cao.
Phan ng c cam ng bi cac tia co bc song dai lien quan vi chlorophyll a
va c thc hien bi quang he thong I. Trong khi o oxy uc giai phong trong trong
cac phan ng do quang he thong II thc hien vi s tham gia cua ca hai chlorophyll va
cac sac to quang hp khac (carotenoid va phycobilin). Hoat ong cua hai loai phan
ng nay lien quan vi cac tia co bc song ngan hn. Tat ca cac te bao quang hp giai

phong oxy eu phai co hai he thong quang hoa, con vi khuan ch co he thong quang
hoa I. Trong qua trnh tien hoa co le he thong quang hoa I xuat hien trc.
He thong quang hoa II xuat hien sau cu ng vi kha nang s dung nc lam chat
kh va do o dan en giai phong oxy.
Moi he thong quang hoa co tap oan sac to ac trng trong mang thylacoid cua luc lap.
n v tiep nhan anh sang cua he thong quang hoa I cha gan 200 phan t chlorophyll a va
gan 50 phan t carotenoid, trong o mot phan t chlorophyll a duy nhat thc hien chc nang
cua bay exiton (tc sa c to trung tam, ky hie u la P 700 ). He tho ng quang ho a II ch a khoang
200 phan t chlorophyll a va 200 phan t chlorophyll b, c hoac d, tuy thuoc tng loai c the.
Ngoai ra mot so tao con co phycobilin. Trung tam cua he thong quang hoa nay cung la mot
phan t chlorophyll a ky hieu la P 680 .
Ca hai loai phan ng sang eu dan en tong hp ATP. S khac nhau gia chung the
hien ac iem cua dong ien t. Tren c s cua dong ien t ma mot trong hai c che tong
h p ATP c goi la quang phosphoryl-ho a co tnh chu ky, con c che kia c goi la quang
phosphoryl-hoa khong co tnh chu ky.
Quang phosphoryl-hoa co tnh chu ky.
Quang phosphoryl hoa co
tnh chu ky, nh mo ta trong hnh
IX.14, lien quan vi dong ien t
kn. Hoat ong cua dong ien t nay
ch lien quan vi he thong quang
hoa I. Sac to trung tam cua he thong
(P700) sau khi b kch ong bi
nang lng cua photon do cac phan
t sac to anten chuyen en, the kh
tieu chuan cua no c nang len +
0,4V len - 0,6V, do o tr thanh mot
chat kh kha manh. Phan t chat
nhan au tien e no chuyen ien t
giau nang lng en la mot enzyme
cha sat va lu huynh (FeS). Chat
nay b kh va P700 b oxy-hoa
trong qua trnh van chuyen ien t
nay. ien t sau o tiep tuc c
van chuyen qua mot loat enzyme
tren mang thylacoid cua luc lap.
Cuoi cung ien t c chuyen cho
plastocyanin (PC), tai o no ch e
c lap vao cho trong va chu trnh
c khep kn.
Hnh IX.14. Quang phosphoryl hoa co
tnh chu ky.

Khi ien t giau nang lng ri khoi phan t P700, no co mc nang lng cao;
khi quay ve, no tr nen ngheo nang lng. S giam sut nang lng nay xay ra dan
dan. Khi ien t chuyen t chat van chuyen nay en chat van chuyen khac trong
chuoi, no giai phong mot phan nang lng, lam cho gradient nang lng giam tng
bc t trang thai kch ong xuong trang thai c ban, moi bc giam mot phan vi
mc o sao cho te bao s dung mot cach thuan li. Mot phan trong so nang lng nay
c dung e tong hp ATP theo c che hoa tham thau.
Ve kha canh nang lng, c che phosphoryl-hoa co tnh chu ky co hieu qua khong ln.
Trong so khoang 25KCal/mol nang lng thu nhan c do kch ong P700 ch co phan nang
lng giai phong khi ien t i t plastoquinon (PQ) en cytochrome f (Cyt f) vi mc
3,4KCal/mol la thc s co ch cho te bao. e khac phuc tnh hieu qua kem nay, trong qua
trnh tien hoa a xuat hien mot c che quang phosphoryl-hoa khac - quang phosphoryl-hoa
khong co tnh chu ky.
Quang phosphoryl-hoa kho ng co tnh chu ky.
Quang phosphoryl-hoa khong co tnh chu ky loi cuon s tham gia cua ca hai he
thong quang hoa. C che cua no c mo ta trong hnh IX.15. Giong nh quang
phosphoryl-hoa co tnh chu ky, ay qua trnh cung c bat au khi mot photon tac
ong len phan t chlorophyll anten va sau o kch ong P700. Chat nhan ien t t
P700 la FeS va feredoxin (Fd). Nhng ay ket thuc s giong nhau gia hai con
ng. Thay v tiep
tuc ri xuong theo
mach kn, ien t c
chuyen t Fd cho mot
enzyme phu thuoc
flavin (FP). Sau o FP
chuyen ien t nhan
c cho mot chat
nhan quang trong la
NADP
+
e kh chat
nay thanh NADP.H.
Cac phan t anten va
P700 cua trung tam
phan ng cung vi
chuoi van chuyen ien
t t FeS en FP la
thanh phan cau tao cua
he thong quang hoa I.
Cac phan t
NADP.H sau o c
dung e kh CO
2
trong
Hnh IX.15. Quang phosphoryl hoa khong co tnh chu ky.

pha toi cua quang hp
e tong hp glucose va
cac hp chat hu c khac. Nh vay, ay dong ien t i t P 700 qua chuoi van
chuyen en NADP
+
roi cuoi cung en glucose. Mot phan nang lng giai phong trong
chuoi van chuyen ien t nay cung c s dung e tong hp ATP.
Do cac ien t giau nang lng cua P700 b gi lai trong NADP.H va cuoi cung
a vao glucid, nen sac to nay b thieu hut ien t. Nhng "ho ien t" nay c lap
mot cach gian tiep nh cac ien t lay t nc nh hoat ong cua quang he thong II.
Khi anh sang vi bc song tng ng tac ong len mot sac to anten cua quang
he thong II, nang lng c chuyen cho sac to trung tam P680. Phan t nay en lt
mnh se nhng ien t giau nang lng cho mot chat nhan ky hieu la Q. Sau o Q
chuyen ien t cho mot chuoi chat nhan giong nh trong phosphoryl-hoa co tnh chu
ky, tng bc lam giam gradient nang lng en mc cua "ho ien t" trong P700 cua
he thong quang hoa I. Khi ien t di chuyen t tren xuong doc theo chuoi, mot phan
nang lng giai phong cung c te bao s dung e tong hp ATP. Nh vay, cac "ho
ien t" xuat hien trong bien co anh sang th nhat c lap bang cac ien t di chuyen
t he thong quang hoa II en nh bien co anh sang th hai. Luc nay, cac ien t t
nc bat au thc hien vai tro cua mnh. P680 do mat ien t nen tr thanh mot chat
oxy-hoa manh. Nh s ho tr cua mot phc he enzyme ky hieu la Z, no thc hien s
cuon hut ien t t nc, e lai ang sau cac ion H
+
va oxy phan t:
2H
2
O 4e
-
+ 4H
+
+ O
2

en vi P 680
Nh phan ng nay, oxy c giai phong t nc nh mot san pham phu. Cac ion
H
+
cung ong vai tro quan trong cua mnh trong viec tong hp ATP.
4. C s cau truc cua quang phosphoryl-hoa.
Luc lap cung c to chc tng t nh ty the e co the tao nen va duy tr
gradient hoa tham thau, lam ong lc cho sinh tong hp ATP va cac qua trnh trao oi
chat khac. Cac sac to anten cung vi cac trung tam phan ng (QHT I va QHT II), cac
phan t cua chuoi van chuyen ie n t va he enzyme tong hp ATP (CF
0
va CF
1
) eu
nam tren mang thylacoid cua luc lap. Giong nh mang trong cua ty the, mang
thylacoid tao ieu kien e phat sinh gradient ien hoa, cung cap nang lng cho tong
hp ATP. Tuy nhien, khac vi ty the, trong luc lap ion H
+
c tch luy trong ngan noi
chat cua thylacoid, con ngan ngoai, tc noi chat cua luc lap (stroma) th tch ien
am.
Cac qua trnh van chuyen ien t, hnh thanh gradient ien hoa va s dung
gradient nay e tong hp ATP c mo ta trong hnh IX.16.
Qua trnh quang hp c bat au khi photon c hap thu bi phc he cac sac
to anten cua he thong quang hoa I. Nhng e thuan tien, ta bat au xem xet t s kch

ong cua mot phoron len he thong quang hoa II. Nang lng kch ong c chuyen
en phan t sac to trung tam P 680 (nam trong QHT II) va lam cho no tch luy nang
lng. P680 chuyen ien t giau nang lng cho chat nhan Q. Ho ien t trong P 680
c lap nh he enzyme von lam nhiem vu phan giai nc, tch luy mot ion H
+
ben
trong thylacoid khi moi ien t c chuyen cho P 680. ien t giau nang lng ban
au c chuyen en Q, mot phan t di ong qua lai gia hai pha cua mang
thylacoid. Mot phan nang lng cua ien t c dung ay e di chuyen mot ion H
+
t stroma vao trong thylacoid, lam cho tat ca co hai ion H
+
c tang cng cho noi
chat cua thylacoid.

Hnh IX.16. C s cau truc cua phosphoryl hoa quang hp

ien t c chuyen tiep tuc cho phc he cytochrom bf va sau o cho PC, tai o
no ch e lap vao cho trong trong P700 (nam trong QHT II). Khi mot photon c hap
thu bi mot phan t anten trong quang he thong I va kch ong mot ien t, nang
lng kch ong c chuyen cho phan t P700, lam cho no tch c nang lng.
ien t giau nang lng cua P700 nhanh chong c chuyen cho chat nhan Fd. Ho
ien t xuat hien trong P700 bay gi co the c lap bi ien t he thong quang hoa II
ang ch trong PC.
Trong khi o ien t giau nang lng trong he thong quang hoa I chuyen en Fd,
t o co the i theo hai hng. Trong ieu kien bnh thng ien t nay c dung e
kh NADP
+
thong qua flavoprotein (FP) von co mat trong mang. Trong qua trnh nay
tieu thu mot ion H
+
cua stroma e tao ra NADP.H.
Dong ien t a mo ta dan en hai ket qua: Mot la tao ra phan t NADP.H giau
nang lng e tham gia tong hp glucid, mat khac tao ra gradient ien hoa e sau o

c dung cho tong hp ATP. Cung nh trong ty the, mang thylacoid cha phc he
enzyme CF
1
co kha nang s dung nang lng cua gradient ien hoa e phosphoryl-
hoa ADP.
5. Co nh CO
2
trong pha toi cua quang hp.
a/ Chu trnh Calvin.
Sau nhieu nam nghien cu (1946-1961) Melvin Calvin thuoc trng ai hoc
Berkeley (Hoa ky) vi cong cu chu yeu la
14
O
2
a tm ra mot trat t cac phan ng co
nh CO
2
trong quang hp ma ngay nay chung ta goi la chu trnh Calvin. Chu trnh nay
con c goi la con ng C
3
do san pham c hnh thanh au tien la ot hp chat 3
carbon acid 3-phosphoglyceric. Toan bo chu trnh bao gom cac phan ng sau ay:
E
1
1/ 6CO
2
+ Ribuloso-1,5-di (P) 12 Acid 3-Phosphoglyceric
E
2
2/ 12 Acid 3-(P)-glyceric + 12ATP 12 Acid 1,3-Di(P)-glyceric + 12 ADP
E
3
3/ 12 Acid 1,3-Di(P)-glyceric + 12NADP.H + 12 H
+

12 Glyceraldehyde-3-(P) + 12NADP
+
+ 12P
vc
E
4
4/ 5 Glyceraldehyde-3-(P) 5 Dioxyacetone-phosphate
E
5
5/ 3 Glyceraldehyde-3-(P) + 3 Dioxyacetone-phosphate
3 Fructoso-1,6-di(P)
E
6
6/ 3 Fructoso-1,6-di(P) 3 Fructoso-6-(P) + 3P
vc
E
7
7/ Fructoso-6-(P) Glucoso-6-(P)
E
8
8/ Glucoso-6-(P) + H
2
O Glucose + Pvc
E
9
9/ 2 Fructoso-6-(P) + 2 Glyceraldehyde-3-(P)
2 Xyluloso-5-(P) + 2 Erytroso-4-(P)
E
5
10/ 2 Erytroso-4-(P)+ 2 Dioxyacetone-phosphate
2 Cedoheptuloso-1,7-di(P)
E
10
11/ 2 Cedoheptuloso-1,7-di(P) + 2H
2
O
2 Cedoheptuloso-7-(P) + 2 P
vc
E
9
12/ 2 Cedoheptuloso-7-(P) + 2 Glyceraldehyde-3-(P)
2 Riboso-5-(P) + 2 Xyluloso-5-(P)
E
11
13/ 2 Riboso-5-(P) 2 Ribuloso-5-(P)
E
12

14/ 4 Xyluloso-5-(P) 4 Ribuloso-5-(P)
E
13
15/ 6 Ribuloso-5-(P) + 6 ATP 6 Ribuloso-1,5-di(P) + 6 ADP
Enzyme xuc tac chuoi phan ng tren ay bao gom:
E
1
: Ribulosophosphate carboxylase (Ribulosophosphate carboxydismutase)
E
2
: Phosphoglycerate kinase
E
3
: Glyceraldehydephosphate dehydrogenase
E
4
: Triosophosphate isomerase
E
5
: Aldolase
E
6
: Fructosodiphosphatase
E
7
: Hexosophosphate isomerase
E
8
: Glucoso-6-phosphatase
E
9
: Trancetolase
E
10
: Phosphatase
E
11
: Isomerase
E
12
: Epimerase
E
13
: Phosphoribokinase
Phng trnh tong quat cua toan bo chu trnh la:
6CO
2
+ 18ATP + 12NADP.H + 12H
+
Hexose + 18ADP +
+ 18P
vc
+ 12NADP
+
+ 6O
2
b/ Chu trnh Hatch Slack
Vao nhng nam 1960 hai nha bac hoc M.D. Hatch (ngi Uc) va Slack (ngi
Anh) a phat hien mot c che sinh hoa cay ma va mot so thc vat nhiet i khac,
trong o san pham au tien cua qua trnh kh CO
2
la mot hp chat 4 carbon acid
oxaloacetic. Chu trnh nay v vay c goi la con ng C
4
trong quang hp. No co
dang nh mo ta trong hnh IX.17.
nhng thc vat co con ng C
4
(goi tat la cay C
4
) hoat tnh enzyme PEP-
carboxylase enzyme then chot cua chu trnh C
4
cao gap 20 lan so vi cay C
3
. Li
the cua cay C
4
la cho bang cach tong hp oxaloacetate va malate chung d tr c
CO
2
trong te bao, nh o chung co the thc hien c quang hp trong luc kh khong
ong do nhiet o moi trng qua cao e han che thoat hi nc. V ly do nay ma thc
vat C
4
thng co tnh chu han cao. Mot ne t u viet khac cua thc vat C
4
la han che
c tac hai cua quang ho hap ma thc vat C
3
thng thieu ot mat khoang 50% san
pham trung gian cua chu trnh Calvin.
Nguyen nhan cua hien tng quang ho hap la do ribulosophosphate
carboxylase von xuc tac phan ng au tien cua chu trnh Calvin trong ieu kien ham
lng CO
2
thap co the chuyen sang xuc tac phan ng oxy hoa ribulosophosphate.

Nhiet o cao cua moi trng cung tao ieu kien thuan li cho qua trnh oxy hoa nay.
S oxy hoa nay khong dan en hnh thanh ATP nen cac chat hu c b thieu ot mot
cach vo ch.
thc vat C
4
nh ac iem giai phau cua la a gop phan ngan nga c tac
hai cua quang ho hap. thc vat C
3
cac te bao bao boc xung quanh bo mach khong
cha luc lap, do o quang hp ch xay ra trong cac te bao tht la. Hn the na, te bao
tht la khong tiep xuc chat che vi bo mach. Trong khi o thc vat C
4
cac te bao bao
boc xung quanh bo mach rat giau luc lap, ong thi te bao tht la (cung cha luc lap)
co xu hng bao boc xung quanh cac te bao bao boc bo mach. ang chu y la trong hai
loai te bao nay cau truc cua luc lap khac nhau va hai loai luc lap thc hien hai chc
nang khac nhau: trong te bao tht la chu yeu xay ra chu trnh C
4
e tch luy CO
2
. Sau
o san pham cua chu trnh C
4
c a vao te bao bao boc bo mach, tai o chung b
decarboxyl hoa e giai phong CO
2
phuc vu cho chu trnh Calvin. Nh vay, nh chu
trnh C
4
ma nong o CO
2
trong te bao bao boc bo mach luon mc cao, lam cho oxy
khong the canh tranh vi no tai trung tam hoat ong cua enzyme RDP- carboxylase.

E
5

CH
2
CO
2
COOH COOH CH
3
C-O~(P) Mg
2+
CH
2
NADP.H + H
+
CH
2
NADP
+
C=O
COOH E
1
C=O E
2
CHOH E
3
COOH
COOH COOH
Phosphoenolpyruvate Oxaloacetate Malate
Pyruvate
E
4

CO
2
E
1
: PEP-Carboxylase; E
2
: Malate dehydrogenase;
E
3
: Malicoenzyme; E
4
: PEP-Carboxykinase;
E
5
: Phosphopyruvate synthase CHU TRNH CALVIN
Hnh IX.17 Chu trnh Hatch Slack
c/ Con ng carbon thc vat CAM.
Ngoai cac con ng C
3
va C
4
, mot so thc vat mong nc nh da, xng
rong va nhieu cay khac song vung sa mac, co ten goi chung la thc vat CAM
(Crassulacean Acid Metabolism), con co mot con ng co nh CO
2
khac. Con ng
nay cung cho phep thc vat thch nghi vi ieu kien kho nong keo dai. Tuy nhien, neu
thc vat C
4
cac c che co nh CO
2
khac nhau c phan biet ve mat khong gian, th

nhom thc vat nay c phan biet ve mat thi gian. Qua trnh carboxyl-hoa s cap
xay ra vao ban em, trong o acid phosphoenolpyruvic ket hp vi CO
2
e tao ra acid
malic. Vao ban ngay, trong ieu kien kh khong ong, CO
2
c giai phong e tham
gia chu trnh Calvin.

CHNG 5. LIPID

LIpId Ia moI nIom Iop cIaI Iuu co nguon goc sInI vaI, co InI k
nuoc nIung Ioa Ian de dang Irong cac dung moI kIong pIan cuc. NIom
Iop cIaI nay Ia moI Irong nIung IIanI pIan cInI cua Ie bao va mo dong
IIuc vaI. MoI so Irong cIung duoc dung Iam cIaI dInI duong du Iru, moI
so kIac IIam gIa Irong vIec kIen Iao cac co quan Iu cua Ie bao, dac bIeI Ia
cac Ie IIong mang, dong IIoI IIam gIa Irong vIec dIeu Ioa cac qua IrnI
van cIuyen vaI cIaI qua mang va nIIeu qua IrnI quan Irong kIac .
Phan ln lipid la dan xuat cua acid beo, hnh thanh khi cac nhom carboxyl lien
ket ester vi mot ru a chc hoac n chc. Ngoai ra, trong phan t cua nhieu loai
lipid khac nhau con cha cac nhom ion hoa hoac phan cc nh base nit, phosphate,
saccharide, aminoacid v.v... Mot so hp chat tuy khong phai la dan xuat cua acid beo
nhng do co tnh k nc nen cung c xep vao nhom lipid. o la sterol va cac hp
chat steroid, sac to quang hp, cac vitamin tan trong chat beo va vai hp chat khac.
Nhng loai lipid ma trong phan t cua chung cha ong thi cac nhom chc
phan cc va khong phan cc thng gap trong cac cau truc mang va tren cac be mat
phan cach khac gia moi trng nc va cac khu vc k nc ben trong te bao.
I. ACID BEO.
Acid beo la thanh phan cau tao cua phan ln lipid. Hn 70 loai acid beo tm thay
trong cac loai te bao khac nhau la nhng mach hydro carbon no hoac khong no cha
mot nhom carboxyl tan cung. Hau het cac acid beo co so chan nguyen t carbon t 12
en 22, nhng thng gap nhat la nhng acid beo co 16 18 nguyen t carbon. Acid
beo khong no co the cha mot hoac vai lien ket oi. Rat t gap acid beo cha lien ket
ba. Phan ln cac lien ket oi noi C
9
vi C
10
(k hieu la
9
). Cac lien ket oi th hai va
th ba thng nam cach lien ket oi th nhat mot vai nhom methylen (-CH
2
-) ve pha
au CH
3
tan cung. Cac lien ket oi trong phan ln acid beo khong no trong t nhien co
cau hnh cis-, mac du dang nay kem ben vng hn dang trans-. Khi un nong vi mot
so chat xuc tac, dang cis- se chuyen sang dang trans-, ong thi nhiet o nong chay
tang len.
Dang cis- cua acid beo khong no lam cho nhng phan t cha nhieu lien ket oi
c uon cong va co ngan lai. ac iem nay co y ngha sinh hoc quan trong ac biet
oi vi cau truc cua cac loai mang te bao. oi vi acid beo no dang duoi thang thch
hp hn ve mat nang lng, mac du tren nguyen tac chung co the ton tai vo so kieu
cau hnh khac nhau do cac lien ket CC trong mach hydro carbon quay mot cach
hoan toan t do. Kha nay acid beo khong no b han che do cac lien ket oi khong
xoay c. Nhiet o nong chay cua acid beo no cao hn acid beo khong no.

Pho bien nhat trong t nhien la nhng acid beo no va khong no gii thieu trong
hnh III.1.

Trong acid beo khong no oi khi con cha lien ket ba -CC-, mac du rat t gap,
v du acid crepinis trong hat Crepis foetida (thuoc ho Compositae).
Hnh 3.1. Cac acid beo no va khong no pho bien
Hnh III.1. Nhng acid beo pho bien trong t nhien.
Mot so acid beo con
cha nhom OH, v du acid
ricinic trong hat thau dau
(Ricinus communis)
vi khuan con gap cac
acid beo cha chom
xyclopropyonyl, lam cho
chung tr nen co so le nguyen
t carbon, v du acid
lactobaxilis.
Mot so acid beo co so le nguyen t carbon con do chung cha nhom CH
3

mach nhanh tng t nh trong phan t cua cac aminoacid valine, leucine, isoleucine.
Acid beo hau nh khong tan trong nc, nhng muoi Na hoac K cua chung (xa
phong) co the tao nhng chuoi phan t goi la mixen (micella) kha on nh trong nc
do tac dung cua tng tac k nc gia chung, nh o xa phong co tac dung tay ra.

Acid beo khong no de ket hp vi hydro hoac halogen tai cac v tr lien ket oi.
Nh tnh chat nay ngi ta co the bien acid beo khong no thanh acid beo no trong
cong nghe che tao m thc vat hoac xac nh t le lien ket oi trong phan t acid beo
bang cach cho tac dung vi I
2
hoac Cl
2
.
Mot so acid beo khong no (acid linoleic, acid linolenic, acid arachidonic...)
khong c tong hp trong c the ngi va ong vat co vu. V the chung thng c
goi la acid beo khong thay the, hay vitamine F.
Cac acid beo t do thng gap trong t nhien thong thng nam tren cac be mat
phan cach gia lipid va nc, au mang goc COOH cua chung phan li thanh COO
-

va quay ve pha moi trng nc. Tuy nhien, phan ln acid beo lien ket vi cac thanh
phan khac nhau trong cac loai lipid khac nhau bang lien ket ester hay lien ket amid.
Mac du acid beo rat a dang, nhng trong moi loai c the ch co mot vai loai la
co ham lng ang ke. thc vat bac cao chu yeu co mat acid palmitic va va hai acid
beo khong no la acid oleic va acid linoleic. Acid stearic hau nh khong co thc vat,
con cac acid beo C
20
C
24
c gap rat t (tr bieu b cua la). Trong khi o mot so
loai thc vat ac biet lai cha mot so loai acid beo ac biet vi ham lng kha cao, v
du acid crepinic chiem 60% tong so acid beo trong hat Crepis foestida, con acid ricinis
chiem en 90% acid beo cua hat thau dau Ricinus communis.
Trong c the ong vat cung cha acid palmitic va acid oleic, nhng ngoai ra con
co acid stearic vi ham lng kha cao.
vi khuan thng khong co cac acid beo cha nhieu lien ket oi nhng lai
thng gap cac acid beo cha nhom cyclopropionyl va nhom hydroxyl trong thanh
phan cua cac loai lipid khac nhau hoac trang thai t do.
Ham lng acid beo trong cac c quan khac nhau cua cung mot c the thng rat
khac nhau. V du, trong thanh phan lipid cua mang t le acid beo khong no cao hn
trong mo m d tr.
II.CAC ESTER CU A GLYCEROL.
Nhng lipid ma ban chat hoa hoc cua chung la ester cua glycrol (glycerine) bao
gom cac hp chat khac nhau ma ten goi va cau tao cua chung c gii thieu trong
bang III.1.
1.Lipid trung tnh.
Glycerol co the lien ket vi mot, hai hoac ba phan t acid beo e tao ra mono-,
di- hoac triglyceride. Ca ba hp chat nay c goi chung la lipid trung tnh, nhng pho
bien nhat trong t nhien la triglyceride, hay triacylglycerine. o la thanh phan chnh
cua dau va m ong thc vat. Phan ln triglyceride co thanh phan a hp, tc hai hoac
ba loai acid beo khac nhau, trong o co acid beo no va khong no, thng la nhng acid

beo 16 18C. Cac triglyceride n gian, tc ch cha mot loai acid beo, v du triolein
trong dau oliu, rat t gap trong t nhien. Dau thc vat cha nhieu acid beo khong no
nen de nong chay hn m ong vat.

Bang III.1. Cac lipid la ester cua glycerol
H
2
C CH CH
2
O O O
Goc glycerine
FA FA FA Lipid trung tnh
FA FA P Acid phosphatidic
FA FA P O CH
2
CH
2
NH
3
+
Goc choline
Phosphatidylcholine
(Leucitine)
FA FA P O CH
2
CH
+
N (CH
3
)
3
Goc ethanolamine
Phosphatidylethanolamine
(Cephaline)
FA FA P O CH
2
CH NH
3
+
COO
-

Goc serine

Phosphatidylserine
FA FA P OH OH

O
HO OH
OH
Goc inositol

Phosphatidylinosit(ol)
(Inositide)
FA FA P O OH OH
H
2
C CH CH
2
Goc glycerine

Phosphatidylglycerine
FA FA P O OH
H
2
C CH CH
2
O
P
O
H
2
C CH CH
2
O O
FA FA

Diphosphatidylglycerine
(Cardiolipin)
HC FA P O CH
2
CH NH
3
+
HC hoac O CH
2
CH
2

+
N (CH
3
)
3

Plasmalogen

R
FA FA Mot hoac hai goc galactose Glycerogalactolipid
FA FA Gal-6-SO
3
-
Glycerosulphate
Triglyceride c goi ten tren c s thanh phan acid beo, v du palmitodiolein,
palmitooleinolinolein v.v... Dau cung nh m thng la hon hp cua nhieu loai
triglyceride khac nhau.
Lipid trung tnh c c the s dung chu yeu
lam chat dinh dng d tr, ch mot phan rat t tham
gia cau truc mang te bao.

Acid beo khong no dc tac dung cua anh sang,
tia ion hoa, oxy khong kh va o am cao rat de b oxy
hoa. Giai oan au cua qua trnh nay la s hnh thanh
peroxyde.

R CH = CH R R CH CH R
O O
Sau o peroxide tiep tuc b phan giai thanh cac aldehyde co mui kho chu:

O O
R CH CH R R + R
O O H H
Qua trnh nay lam cho dau va m giam chat lng. e han che no, cung nh e
de ong goi va van chuyen,ngi ta tien hanh hydrogen hoa cac lien ket oi, lam cho
dau bien thanh m thc vat.
Di tac dung cua cua acid va cua enzyme lipase lipid trung tnh b thuy phan
thanh glycerol va acid beo t do. Neu thuy phan bang kiem se thu c muoi cua acid
beo (xa phong).
Tnh chat cua tng loai dau hoac m thng c s dung bang cac ch so sau
ay:
Ch so iod: la so mg iod ket hp vi 100 gam dau hoac m. No cho phep tnh
c t le acid beo khong no trong dau,m;
Ch soacid: la so mg KOH can e trung hoa toan bo acid beo t do trong 1 gam
dau hoac m. Ch so nay cho phep kiem tra s hnh thanh acid beo t do trong qua
trnh bao quan dau, m;

Hoa sinh hoc
- 122 -

Ch so xa phong hoa: la la so mg KOH can e trung hoa toan bo acid beo t do
va lien ket co trong 1 gam dau hoac m. Trong lng phan t cua triglyceride cang
ln th ch so xa phong hoa cang nho.
2.Phosphatide.
Phosphatide la nhng lipid phc tap co cha acid phosphoric. Tuy nhien, thuat
ng nay thng dung e ch nhng phosphatide la ester cua glycerol, tc
phosphoglyceride, hay glycerophospholipid. ai dien n gian nhat cua nhom lipid
nay la acid phosphatidic. No hnh thanh khi hai nhom OH t do cua glycerol ester
hoa vi acid beo, con nhom OH th ba - vi acid phosphoric. Trong c the acid
phosphatidic c tong hp t -glycerophosphate tc dang hoat ong cua glycerol. Tuy
acid phosphoric ton tai trong te bao dang t do vi ham lng rat thap, nhng la mot
san pham trung gian quan trong cua qua trnh sinh tong hp lipid trung tnh va cac
phosphoglyceride khac. Nhng phospho-glyceride nay hnh thanh khi goc phosphate
cua acid phosphatidic ester hoa vi mot base nit (choline, ethanolamine, serine)
hoac vi mot chat khac co cha chc ru t do (inositol, glycerine v.v...) (bang III.1).

ac iem cau tao va
cha nhom k nc (acid
beo), va cha cac nhom a
nc (goc phosphate, base


nit, glycerine v.v...) cho
phep phosphoglyce-ride
tham gia cac cau truc mang
cua te bao va ieu hoa cac
qua trnh van chuyen vat
chat qua mang.
Trong mo than kinh, tim, gan, trng cua o ng vat co xng song va trong hat cua
thc vat ham lng phosphoglyceride rat cao.
Trongphosphatidylcholine (leucitine) goc phosphate cua acid phosphatidic ester
hoa vi choline. Tat ca cac acid beo pho bien trong lipid trung tnh eu co mat trong
phosphatidylcholine. Acid beo no thng lien ket vi nhom OH cua C, con acid beo
khong no vi C.
Phosphatidylethanolamine (cephaline) va phosphatidylserine cung la nhng
phosphoglyceride pho bien. Vi s tham gia cua S-adenosylmethionine (dang hoat
ong cua methionine) phosphatidylethanolamine co the c methyl hoa thanh
phosphatidylcholine, con phosphatidylserine nh enzyme ac hieu co the b
decarboxyl hoa thanh phosphatidylethanolamine.
Phosphatidylinosit, hay inositolphosphatide, cha ru vong inositol. Cac nhom
hoa tr -OH co the nh v xch ao hoac quanh truc, cho phep hnh thanh 9 dang
ong phan khac nhau. Nhng nhom OH nay thng c ester hoa vi acid
phosphoric, tao ra diphosphoinositide, triphosphoinositide va cac dan xuat bac cao
hn.
Phosphatidylglycerine va diphosphatidylglycerine (cardiolipine) cha them cac
phan t glycerine lien ket thong qua goc phosphate.
Trong tat ca cac loai mang sinh hoc ben canh hang loat cac loai lipid khac nhau
ham lng phospholipid thng chiem u the vi t le t 40 en 90% lipid tong so cua
cau truc mang, trong o nhieu hn ca la phosphotidylcholine,
Phosphatidylethanolamine, phosphatidylserine va cardiolipine. Phosphatidylinosit
thng co mat vi ham lng thap hn. Ham lng cardiolipine ac biet cao trong
mang cua vi khuan, ti the va luc lap.
3.Glycerogalactolipid va glycerosulfolipid.
Trong so cac esIer cua gIycerIn con co gIyceroIacIoIIpId (gaIacIosyI
dIgIycerIde) va gIycerosuIIoIIpId.

Hoa sinh hoc
- 124 -

Hai loai lipid nay la thanh phan cau tao quan trong cua mang te bao thc vat noi
chung va mang luc lap noi rieng. Trong galactosyldiglyceride cua luc lap t le acid
linolenic chiem en 96% acid beo tong so.
Can phan biet nhom lipid nay vi glycolipid va sulfolipid (sulfatide) thuoc nhom
sphingolipid se c xet en sau ay.
III. XPHINGOLIPID VA GLYCOLIPID.
Hai nhom lipid nay eu cha mot loai ru amin co ten la sphingosine. Song
gia ru va acid beo gan vi nhau khong phai bang lien ket ester ma bang lien ket
amide. Sphingolipid n gian nhat la ceramide. Neu chc ru bac mot cua
sphingosine c gan them goc phosphorylcholine th se tao ra mot sphingolipid khac
la sphingomyeline.
Co the xem glycolipid la nhng sphingolipid co cha glucid hoac dan xuat cua
glucid. Cau tao cua cac nhom sphingolipid va glycolipid khac nhau c gii thieu
tom tat trong bang III.2.
Bang III.2. Cau tao cua sphingolipid va glycolipid

Can lu y rang, khac vi sphingolipid,
glycolipid khong cha acid phosphoric.
Cung vi phospholipid, sphigolipid va
glycolipid cung gop phan quan trong trong
viec hnh thanh cac cau truc mang, trong
sphingo-myeline thng co ham lng kha
cao. Tuy nhien, loai sphingolipid nay lai
khong co mat trong ti the.

Cac loai glycolipid
khac nhau co nhieu trong
mao, lach va cac c quan
khac cua ong vat.
Ganglioside la thanh phan
quan trong cua mang te
bao than kinh va tham gia
vao viec phuc hoi ien
kch thch cua nao, loai tr
tnh oc cua oc to vi
trung uon van, bach hau
va cac chat oc khac do vi
khuan tiet ra. Trong mang
te bao than kinh con co
nhieu loai cerebroside va
cac ester sulfate cua
chung, tc sulfolipid hay
cereroside sulfatid.
Sphingolipid khong c tm thay trong te bao thc vat. Tuy nhien, trong nam
men, nam moc va mot so thc vat khac co mot nhom lipid tng t sphingolipid ma
thanh phan ru la cerebrine hay con co ten la phytosphingosine.
IV. SA P.
Sap nguon goc sinh hoc la ester cua mot acid beo bac cao, bao hoa hoac khong
bao hoa cha 14 - 36 nguyen t carbon, vi mot ru n chc phan t ln cha 16 -
30 nguyen t carbon. Nh vay, cau tao cua chung gan vi acylglycerine. Nhiet o
nong chay cua sap t 60 en 100
o
C, tc cao hn nhieu so vi triacylglycerol. Sap tao
nen lp bao ve tren da, long ong vat, qua, than, la thc vat va tren lp vo ngoai cua
nhieu loai con trung. Ngoai ra, mot so plankton song trong nc bien sap c s
dung lam nguon nang lng d tr.
Thanh phan chu yeu cua
sap ong la ester cua acid
palmitic vi ru beo 1-
triacontanol. Lanoline, tc sap
long cu, la hon hp cac ester
cua acid beo vi hai loai ru

Hoa sinh hoc
- 126 -

vong la lanosterine va
agnosterine.
Sap thc vat la hon hp cac ester cua acid beo cha 20-24C vi cac ru beo
cha 26-34C. Hon hp nay thng co thanh phan rat ac trng oi vi tng loai cay.
Ngoai ra, trong thanh phan cua sap thc vat con co cac acid beo va ru beo t do
cung vi mot so hydro carbon, chu yeu la cac alcan vi so le nguyen t carbon t 25
en 35.
Chc nang bao ve be mat cua sap lien quan vi tnh khong phan cc cao va tnh
k nc manh cua chung.
Sap c s dung rong rai trong ky nghe dc pham, my pham va nhieu lnh vc
cong nghe khac.
V. STEROL VA STEROID.
Cac hp chat steroid c ac trng bi bo khung cyclopentanoperhydro-
phenantren.
Sterin (hay sterol) la nhng steroid co
cha chc ru. Chc ru nay thng gan tai
v tr C
3
. Con tai v tr C
17
thng gan mot
mach n/hanh cha 8-10 nguyen t carbon.
Phan ln steroid cha hai nhom CH
3
vi ky
hieu C
18
va C
19
.
Cholestanol co the c xem la mot
sterine ien hnh ve phng dien cau truc,
mac du ham lng cua no trong c the ong
vat khong cao.

Dan xuat dehydrogen hoa cua
cholestanol la cholesterine (cholesrerol). No
ong vai tro quan trong trong cau truc va hoat
ong chc nang cua mang te bao ong, thc
vat. Ngoai ra, cholesterine con chiem v tr
hang au trong trao oi steroid. T no te bao
ong vat tong hp nen cac hp chat steroid co
hoat tnh sinh hoc rat quan trong nh acid mat,
cac hormone tuyen thng than (cortisol,
cortiocosterone, androsterone), hormone
tuyen sinh duc c (testosterone,
androstenedione, androsterone) va hormone
sinh duc cai (estrone, estradiol, estradial,
progesterone).

ien hnh cho sterol thc vat la
ergosterol. No c goi la provitamine D
2
, v
se chuyen hoa thanh vitamine D
2
di tac
dung cua tia t ngoai. Nam men co ham lng
ergosterol rat cao nen c dung lam nguyen
lieu e san xuat vitamine D
2
. Ngoai ra, thc
vat con co nhieu sterol khac nh stigmasterol,
cytosterol v.v...
Sterine hau nh khong co mat trong vi
khuan.
VI. SA C TO QUANG HP.
Do co tnh k nc nen cac sac to quang hp cung c xep vao nhom lipid,
hay ung hn, nhom cac chat tan trong lipid. Nhom sac to nay bao gom ba loai:
chlorophyll, caroteneoid va phycobilin. Chung tham gia trong pha sang cua quang hp
thc vat. Ca ba loai sac to nay trong te bao ton tai dang cac phc he
chromoprotein. Chlorophyll va caroteneoid lien ket vi protein tng oi long leo
thong qua cac lien ket yeu; trong khi o phycobilin lien ket cong hoa tr vi protein e
tao nen cac phc he kha ben vng.
1.Chlorophyll.
Chlorophyll la sac to quang hp chu yeu cua thc vat bac cao va tao. Trong te
bao thc vat eukaryote chung la thanh phan cau tao cua luc lap c quan t lam
nhiem vu quang hp. Co 4 loai chlorophyll khac nhau, ky hieu la a, b, c va d.

Hnh III.1. Cong thc chung cua chlorophyll.

Phan t chlorophyll cha 4 goc pyrol lien ket vi nhau, lam thanh nhan
porphyrine; gia nhan la phan t manhe. Cac loai chlorophyll khac nhau phan biet
nhau bi cac nhom chc gan vi nhan porphyrine va trang thai lien ket gia C
7
vi C
8

(hnh III.1 va bang III.3).
Bang III.3. Ban chat hoa hoc cua cac nhom cha R
1
, R
2
, R
3
, R
4
va trang thai lien
ket gia C7 va C8 cac loai chlorophyll khac nhau.

Chlorophyll a co mat trong tat ca cac loai cay xanh. Hn na, trong moi thc vat
ham lng cua no eu cao hn tat ca cac loai chlorophyll khac.
Chlorophyll b cung pho bien gan nh chlorophyll a. No ch khong co tao, tr
Chlorophyceae va Euglenophyceae.
Chlorophyll c c biet co 2 loai: c
1
va c
2
. Ca hai eu co mat trong
Phaeophyceae, Chrysophyceae va Bacillariophyceae.
Chlorophyll d c phat hien mot so loai tao thuoc ho Rhodophyceae.
Tat ca cac loai chlorophyll eu co kha nang hap thu cac tia o cua ang sang mat
tri vi bc song 650 en 680 nm. Kha nang nay lien quan vi he thong lien ket oi
tiep hp cua nhan porphyrine. Trong khi o nh s co mat cua goc phytol k nc ma
phan t chlorophyll nh hng trong he thong mang luc lap theo nguyen tac ac trng
cho moi phan t co tnh phan cc.
S phan bo cua cac loai chlorophyll trong t nhien cung vi caroteneoid va
phycobilin c gii thieu trong bang V.4.
vi khuan quang hp co nhom sac to tng t bacteriochlorophyll tng t
chlorophyll thc vat. Chung cung c cau tao tren c s nhan porphyrine.
2. Caroteneoid
Caroteneoid la cac dan xuat cua isoprene (CH
2
= C CH = CH
2
) bao gom khoang 60
sac
CH
3

to mau vang, da cam va o. Cau truc polyisoprenoid cua nhom sac to nay trong a
so trng hp bao gom 40 nguyen t carbon. Mach polyisoprenoid cua nhieu sac to
c ket thuc hai au bang cac vong ionone, con mot so sac to khac hai au
van trang thai mach h. Phan trung tam cua phan t caroteneoid gom 16 nguyen
t carbon tao thanh mot mach lien ket tiep hp va 4 nhom methyl:

Cac nhom tan cung gan hai au cua mach trung tam nay cac caroteneoid
khac nhau co cau truc khac nhau. V du, licopine hai au co kieu cau truc (1-1), -
carotenee - (1-2), -carotenee (2-3), -carotenee (2-2), epiquineone (2-4),
cantaxantine (4-4)...
Do cIua moI Iuong Ion cac goc ky nuoc nen caroIeneoId IIe IIen InI
ky nuoc raI ro. CIung deu Ia nIung sac Io Ian Irong IIpId va Ioa Ian de
dang Irong cac dung moI Iuu co kIong pIan cuc (benzyne, eIer peIroI...)
cung nIu dung moI Iuu co pIan cuc (eIer eIIyIIc, aceIone...)
Trong luc lap
NIung caroIeneoId cIua oxy duoc goI cIung Ia xanIIopIyII, Irong do o
moI so daI dIen oxy Ion IaI o dang nIom OH, v du Irong IuIeIne,
zeaxanIIne; con o moI so daI dIen kIac o dang epoxIde, v du Irong
vIoIaxanIIne.



Trong Ie bao quang Iop cac caroIeneoId kIac nIau dong vaI Iro Iap
IIu anI sang bo sung cIo cIIoropIyII. ong IIoI, do dac dIem Iap IIu cac
IIa gIau nang Iuong Ion cac IIa mau do von duoc cIIoropIyII Iap IIu manI
nIaI, nen cIung con co cIuc nang bao ve cIIoropIyII va cac cau Iruc kIac
cua Ie bao kIoI b IIIeu doI boI anI sang maI IroI. Cac caroIeneoId cIua
oxy con co IIe IIam gIa cac pIan ung quang pIan nuoc. MoI so
caroIeneoId co nIung cIuc nang Ioan Ioan kIong IIen quan den quang Iop,
v cIung duoc pIaI IIen ca Irong nam va Irong cac mo, co quan kIong IIuc
IIen quang Iop nIu cacI Ioa, IaI pIan, IuI pIan cua moI so IoaI Ioa.
3. Phycobilin.
Phycobilin tro
Rhodophyceae, Cyanophyceae va Cryptophyceae.

Phycobilin cung c cau tao t 4 vong pyrol nhng khong khep kn thanh nhan
porphyrine.
Phycoerythrob
cac phan di n vi nay ch lien ket vi mot trong hai loai sac to e tao ra
phycoerythrin mau o hoac phycocyanin mau lam. Tuy nhien, cung co trng hp ca
hai loai sac to cung co mat trong mot cau t protein nhng mot trong hai loai se chiem
u the.
Trong te bao p
Cung vi caroteneoid, phycobilin c xem la sac to ho tr cho chlorophyll
trong hoat ong quang hp, mac du cung co y kien cho rang chung hoat ong oc lap.
ac iem phan bo cua chung trong t nhien c gii thieu trong bang III.4.
Bang III.4. Phan bo cua cac sac to quang hp chu yeu trong gii thc vat.
C the Chlorophyll Phycobilin Caroteneoid
a b c
1
c
2
d Phycoerythri
n
Phycocyani
n


Hoa sinh hoc
- 131 -

Thc vat bac cao,
dng x, a y
+ + - - - - -
-Carotene,-
Carotene,
Violaxantine,
Neoxantine
Tao Chlorophyceae

+ + - - - - -
-Carotene,
Lutein,
Violaxantine,
Neoxantine
Euglenophyceae + + - - - - -
-Carotene,
Neoxantine,
Diadinoxantine
Phaeophyceae + - + + - - -
-Carotene,
Fucoxantine,
Violaxantine
Chrysophyceae + - + + - - -
-Carotene,
Fucoxantine,
Xanthophyceae + - - - - - -
-Carotene,
Neoxantine,
Diadinoxantine

Baccillariophyceae

+

-

+

+

-

-

-
-Carotene,
Neoxantine,
Diadinoxantine,
Fucoxantine
Cryptophyceae + - - + - + +
-Carotene,-
Carotene,
Alloxantin
Rhodophyceae + - - - + +++ +
-Carotene,-
Carotene,
Lutein, Zeaxantin

Cyanophyceae

+

-

-

-

-

+

+++
-Carotene,
Equinenone
Micoxantine
Zeaxantine
Prochlorophyceae
(chi Prochloron)
+ + - - - - -
-Carotene,
Zeaxantine
Ghi chu: dau + hoac ch s co mat hay vang mat cac sac to.

VII. VITAMIN TAN TRONG LIPID
1.Vitamin A.
Vitamin A co nhieu loai, nhng
quan trong nhat la retinol, tc vitamin A
1
.
no thng ton tai dang ester vi acid -
glucuronic, tch luy chu yeu trong gan ca
va cac ong vat bien khac. Tuy nhien,
nhu cau vitamin A cua ngi con co the
c ap ng qua nhng thc an thc vat
giau carotene. Chat nay con c goi la
provitamin A, v co the b oxy hoa di
Hoa sinh hoc
Khoa Sinh hoc
- 132 -

GS.TS. Mai Xuan Lng
tac dung cua enzyme carotenease trong
gan va niem mac ruot non thanh retinol.
So sanh cau tao phan t cua retinol vi
-, - va -carotene, ta se thay ro khi oxy
hoa mot phan t -carotene se tao ra hai
phan t vitamin A
1
, trong khi o moi phan
t - va -carotene ch cho mot phan t
vitamin A
1
.
Vitamin A
2
(dehydroretinol) la dang oxy hoa cua vitamin A
1
. No c tch luy
trong gan va m ca nc ngot. Ngoai ra, trong gan ca voi con co vitamin A
3
.
Thieu vitamin A ac biet nguy hiem oi vi tre em, v luc mi sinh chung cha
tch luy c vitamin nay. ngi ln retinol co the tch luy trong gan u e c the s
dung trong vong 2 nam. Thieu vitamin A se anh hng en sinh hoat bnh thng cua
ong vat, giam sc e khang oi vi vi trung v no tham gia trong qua trnh sinh tong
hp lisozyme. Tuy nhien, hieu ng pho bien nhat cua viec thieu vitamin A la lam anh
hng en th giac, gay benh kho mat va quang ga do no ong vai tro quan trong trong
c che sinh tong hp sac to cam quang rhodopsin.
Tha vitamin A cung gay ra hang loat cac benh nguy hiem, v du benh don
xng.
2.Vitamin D.
Vitamin D (calciferol) la
mot nhom vitamin chong benh
coi xng. Co 7 loai vitamin D,
ky hieu t 1 en 7, nhng quan
trong nhat la vitamin D
2

(ergocalciferol) va vitamin D
3

(cholecalciferol) hnh thanh t
cac dang tien than tng ng
ergosterol va 7-
dehydrocholesterol.
Provitamin D
3
, tc 7-
dehydrocholesterol, co mat trong
lp m di da cua ngi. Di
tac dung cua tia t ngoai trong
anh sang mat tri no se bien
thanh cholecalciferol (vitamin
D
3
) vi so lng u e tieu dung
hang ngay (7-12g). tre em
nhu cau cao hn (12-25g). Khi
thieu vitamin D can c bo sung
Hoa sinh hoc
- 133 -

bang nam men (trong o cha
nhieu ergosterol, tc provitamin
D
2
), dau ca va mot so thc an
giau vitamin D khac.
Trong c the ngi va ong vat vitamin D ieu hoa qua trnh hap thu Ca
2+
trong
ng ruot, ieu hoa noi can bang Ca
2+
trong mau va trao oi caclcium, phosphore
trong qua trnh tao xng. Phan ln chc nang khong phai do chnh vitamin D thc
hien, ma la do cac san pham oxy hoa cua no: 25-hydrocalciferol, 1,25-dihydro-
cholecalciferol, 1-dihydro-cholecalciferol v.v... Nhng hp chat nay co hoat tnh
sinh hoc cao hn nhieu so vi chnh cac vitamin D san sinh ra chung va the hien tat
ca cac tnh chat cua hormone steroid.
3. Vitamin E.
Vitamin E (tocoferol) co nhieu loai, pho bien nhat la -, -, va -tocoferol.
Nhom vitamin nay ch c tong hp thc vat. Chung tch luy chu yeu trong hat
(lua, lua my), trong dau (ngo, bong, au tng, hng dng...) va trong mot so rau
xanh.
ong vat thieu vitamin E khong nhng
anh hng en kha nang sinh san ma con lam
ton thng c tim va cac c khac, he tuan hoan
va he than kinh. C che tac dung cua chung
khong nhng lien quan vi tac dung chong oxy
hoa e ngan nga qua trnh oxy hoa cac acid
beo khong no trong cac
cau truc mang ma con lien quan vi sinh tong hp enzyme, ac biet la he
enzyme xuc tac qua trnh tong hp hem.
Nhu cau vitamin E ngi vao khoang gan 5mg tre em va 10-15mg ngi
ln va nhieu hn mot t phu n co mang va cho con bu. Nhu cau nay de dang c
ap ng qua che o dinh dng bnh thng, v vay ngi t xay ra tnh trang thieu
vitamin E.
4.Vitamin K.
Vitamin K la mot nhom gom nhieu vitamin co tac dung lam ong mau. Loai
vitamin K c phat hien au tien la phyloquinone ma ngay nay c goi la vitamin
K
1
. Sau o t bot ca tach c nhom vitamin K
2
(menaquinone) co mach nhanh vi
chieu dai khac khau bi so lng n v isoprene.
Vitamin K can cho ngi e
qua trnh ong mau xay ra bnh
thng hoac e tang cng toc o
ong mau khi can thiet. Chung
tham gia vao qua trnh ong mau
mot cach trc tiep, hoac gian tiep
thong qua tong hp protrombine
va cac yeu to ong mau khac.
Ngoai ra, menaquinone con
tham gia mot so qua trnh oxy hoa
kh, ac biet trong quang hp,
phosphoryl hoa oxy hoa, oxy hoa
acid dihydroorotic thanh acid
orotic.
Ngoai vitamin K t nhien, (K
1
va K
2
), con co nhieu chat tng ong c tong
hp bang phng phap hoa hoc song lai co hoat tnh sinh hoc cao hn. Chung c ky
hieu t K
3
en K
7
.
Vitamin K co nhieu trong rau xanh. Ngoai ra, chung con c vi sinh vat ng
ruot tiet ra vi mot lng ang ke. V vay, thieu vitamin K thng ch xay ra vi tre
em mi sinh. Thieu vitamin K ngi ln chu yeu do lam dung sulfamide va cac chat
khang sinh, dan en c che cac vi sinh vat ng ruot co ch cho qua trnh tieu hoa
thc an.
3. Ubiquinone va plastoquinone.
Ubiquinone con uc goi la vitamin Q, hay coenzyme Q. o la mot nhom dan
xuat cua benzoquinone co mach ben cha 6-10 n v isoprene (ky hieu la vitamin Q
6

Q
10
). Chung c tong hp trong c the ong vat, thc vat va vi sinh vat, la mot
trong nhng vitamin rat pho bien trong t nhien. Nh cau truc mang tnh k nc nen
chung tap trung mang trong cua ti the, vi the, bo may Golji, cung vi vitamin K
2

tham gia vao hoat ong ho hap cua c the bac cao va vi sinh vat, am bao viec hap
thu oxy, van chuyen ien t va phosphoryl hoa oxy hoa trong ti the.
Plastoquinone vi
cau tao tng t
ubiquinone co vai tro
quan trong trong quang
hp vi t cach mot
chat van chuyen ien
t.
Vitamin Q con co
vai tro quan trong trong
viec on nh cau truc
mang, trong
o, tng t nh tocoferol nhng thng co hieu suat cao hn, chung ngan nga qua
trnh oxy hoa cac acid beo khong no.

Trong ieu kien bnh thng ubiquinone c tong hp trong c the ngi u
cho nhu cau, nhng khi oi protein hoac oi nang lng tre em xuat hien trieu chng
thieu mau hoac ton thng nao tuy. Vitamin nay cung can cho s phat trien bnh
thng cua phoi do kch thch s hnh thanh hong cau va c s dung rong rai trong y
hoc e kch thch ti the c tim khi b benh tim mach va loan dinh dng c. Ngc lai,
cac benh nhan ung th co nong o vitamin Q cao ngi ta thng dung cac chat
khang sinh antraxyline von co tac dung khang vitamin Q e ieu tr.
VIII. PHAN GIAI LIPID
1.Phan giai lipid trung tnh.
Lipid chiem t 10 en 20% trong lng c the ong vat co vu, trong o thanh
phan chu yeu la lipid trung tnh (triacylglycerol). Loai lipid nay co mat trong tat ca
cac c quan, ac biet la trong cac mo d tr. Trong qua trnh tien hoa cua sinh gii no
a to ra thch hp vi chc nang d tr nang lng, v trong moi phan t cua no cha
en 3 goc acid beo vi mc o kh cao, do o khi b oxy-hoa, nhng acid beo nay san
sinh ra nhieu nang lng hn bat ky mot nhom hp chat nao khac. V du mc nang
lng d tr de huy ong cua lipid trong c the con ngi cao hn gap 100 lan so vi
glucid.
Thanh phan cua lipid d tr bien ong cac loai khac nhau, song trong pham vi
moi loai th thng kha giong nhau. Hn 99% lipid d tr tch luy trong cac mo cua
ngi la triacylglycerol, bat ke no c tch luy au. Noi chung, lipid d tr giau
acid beo no hn lipid trong gan. Cang giau acid beo no th khi b oxy-hoa no cang san
sinh nhieu nang lng.
Triacylglycerol khong ch co chc nang cung cap nang lng cho cac qua trnh
hoat ong song ma con cung cap nguyen lieu e tong hp hang loat cac hp chat khac
nhau trong te bao.
Di tac dung cua enzyme lipase, triacylglycerol b thuy phan thanh glycerol
va acid beo.
Glycerol sau o c loi cuon vao qua trnh glycolys sau khi c hoat hoa
thanh glycerol-3-phosphate nh glycerokinase.
Glycerol + ATP Glycerol-3-phosphate + ADP.
Sau o glycerol-3-phosphate b oxy hoa thanh glyceraldehyde-3-phosphate. S
chuyen hoa tiep theo co the xay ra theo hai hng: tiep tuc b oxy hoa theo con ng

glycolis va chu trnh Crebs thanh CO
2
va H
2
O, hoac bang cac phan ng ngc vi
glycolis e tong hp glucose va cac monosaccharide khac. Trong khi o acid beo b
oxy-hoa bang cac con ng khac nhau ma chung ta se lan lt tm hieu sau ay.
2.Oxy-hoa acid beo.
a/ -Oxy-hoa acid beo no co so nguyen t carbon chan.
Qua trnh -oxy-hoa c bat au bang mot phan ng chuan b, trong o acid beo
c chuyen thanh dang hoa tan trong nc la acyl-CoA, trong o cac nguyen t
hydro - cua goc acid c hoat hoa:
O O
R-CH
2
-CH
2
-C-OH + ATP + CoA-SH R-CH
2
-CH
2
-C-S-CoA + AMP + PP
i

Phan ng nay c xuc tac bi nhom enzyme hoat hoa acid beo gom acetate
thiolase va cac acyl-CoA synthetase. Nhom enzyme nay bao gom t nhat hai loai: loai
th nhat ac hieu oi vi nhng chuoi hydrocarbon co chieu dai trung bnh (4-12
nguyen t carbon); loai th hai ac hieu cho nhng chuoi dai hn. Ngoai ra, trong ty
the con co mot loai acyl-CoA synthetase hoat ong vi s tham gia cua GTP, nhng
khac vi cac enzyme hoat hoa khac, trong phan ng hai phan t GTP lan lt b phan
giai thanh 2 GDP va 2 P
i
.
Mot trong nhng yeu to xac nh toc o oxy-hoa acid beo la toc o xuyen tham
cua chung vao ty the. Trong khi mot so acid beo (khoang 30% acid beo tong so) t
chung co the xuyen tham vao ty the va trong matrix cua bao quan nay se c hoat
hoa thanh acyl-CoA, th phan ln acid beo co mach carbon dai hn khong chui qua
c mang trong cua ty the va do o can phai c hoat hoa ngay sau khi xuyen qua
c mang ngoai cua ty the. Sau o nhng acyl-CoA nay se di chuyen qua lp mang
trong cua ty the nh lien ket tam thi vi mot chat van chuyen la carnitine (-
trimethyl-amino--oxybutyrate).
(CH
3
)
3
N
+
-CH
2
-CH-CH
2
-COOH
OH
Carnitine
Carnitine co mat hau nh trong moi c the va trong tat ca cac mo ong thc
vat. Nh mot acyl transferase ac hieu, goc acyl cua acyl-CoA c chuyen cho nhom
-OH cua carnitine, tao nen acylcarnitine:
O O H N
+
(CH
3
)
3

R-C-S-CoA R-CO O
-

C
CarnitinCoA-SH CoA-SH (Acylcarnitin) O
Ngi ta cho rang acyl-carnitine de xuyen qua mang hn acyl-CoA v cac ien
tch am va dng nam gan nhau hn va de trung hoa nhau. Sau khi i vao matrix cua

ty the va trc khi bat au qua trnh -oxy-hoa, nhom acyl lai c chuyen ve cho
CoA-SH.
Qua trnh -oxy-hoa acid beo no c thc hien qua 4 phan ng ke tiep sau
ay:

Acyl-CoA- dehydrogenase
1/ R-CH
2
-CH
2
-CH
2
-CO-S-CoA + FAD
R-CH
2
-CH=CH-CO-S-CoA + FAD.H
2

Enoyl-CoA- dehydratase
2/ R-CH
2
-CH=CH-CO-S-CoA + H
2
O
R-CH
2
CHOH - CH
2
- CO-S-CoA
-
Hydroxyacyl-CoA- dehydrogenase
3/ R-CH
2
-CHOH-CH
2
-CO-S-CoA + NAD
+

R-CH
2
-CO-CH
2
-CO-S-CoA + NAD.H + H
+
Acyl-CoA-thiolase
4/ R-CH
2
-CO-CH
2
-CO-S-CoA + CoA-SH
R-CH
2
-CO- S-CoA + CH
3
-CO~SCoA

Trc tien, trong phan ng (1), acyl-CoA b oxy-hoa vi s xuc tac cua acyl-
CoA dehydrogenase ma coenzyme la FAD. Trong phan ng nay cac nguyen t H tai
cac v tr - va - c FAD lay i. Ket qua la hnh thanh mot dan xuat khong no
enoyl-CoA. Tiep theo, trong phan ng (2) enoyl-CoA b hydrate hoa, trong o nhom -
OH c gan tai v tr -, con nguyen t hydro - tai v tr -. Ket qua tao ra L--
oxyacyl-CoA. Trong phan ng (3) chc ru tai v tr - b oxy-hoa bi -oxyacyl-
CoA dehydrogenase vi coenzyme la NAD
+
, dan en s hnh thanh -cetoacyl-CoA.
Cuoi cung, trong phan ng (4) vi s xuc tac cua acyl-CoA-thiolase va mot phan t
CoA-SH t do mach cetoacyl-CoA b cat gia C- va C- e tao ra acetyl-CoA va
mot acyl-CoA mi ngan hn acyl-CoA ban au 2 nguyen t carbon. No lai se tiep tuc
b oxy-hoa theo trat t 4 phan ng tren e tao ra phan t acetyl-CoA th hai. Qua trnh
nay se tiep tuc cho en khi toan bo phan t acyl-CoA vi 2n nguyen t carbon b phan
giai thanh n phan t acetyl-CoA.
Qua trat t cac phan ng tren ta thay cung vi viec rut ngan mot n v acetyl-
CoA (C
2
) t phan t acid beo c che -oxy-hoa cho phep tao ra mot phan t NAD.H
va mot phan t FAD.H
2
. Mot phan t acetyl-CoA trong chu trnh acid tricarboxylic se
tao ra them 1 ATP, 3 NAD.H va 1 FAD.H
2
. Trong chuoi ho hap toan bo so coenzyme
kh nay se tao ra 16 phan t ATP. Nh vay, 17 phan t ATP hnh thanh khi mot n
v C
2
cua acid beo b oxy-hoa hoan toan thanh kh carbonic va nc. Mach acid beo
cang dai th trong -oxy-hoa cang tao ra nhieu acetyl-CoA, va do o khi b oxy-hoa
hoan toan se san sinh ra cang nhieu nang lng hn. Tuy nhien, acetyl-CoA sau khi

hnh thanh co the khong b oxy-hoa tiep tuc theo chu trnh acid tricarboxylic ma c
dung cho cac muc ch sinh tong hp theo trat t cac phan ng cua chu trnh
glyoxilate.
b/ Oxy-hoa acid beo no co so nguyen t carbon le.
Phan ln acid beo cha trong cac mo cua c the ngi, ong vat va thc vat co
so chan nguyen t carbon va mach carbon khong phan nhanh. Tuy nhien, trong mot so
vi sinh vat va trong sap thng gap cac acid beo co cau truc phan nhanh, chu yeu bi
cac nhom CH
3
(hnh 10.1). Neu mc o phan nhanh khong nhieu va moi iem
phan nhanh
eu nam tai v tr chan (tnh t
au carboxyl) th qua trnh -oxy-
hoa xay ra bnh thng. Khi mach
acid beo b phan huy, ben canh
acetyl-CoA con tao ra propionyl-
CoA. Neu cac nhom methyl nam
tai cac v tr le th qua trnh -
oxy-hoa b phong toa sau khi thc
hien phan ng acyl-CoA dehydro-
genase. Trong trng hp nay
neu nhom methyl cua mach
nhanh nam sat au tan cung oi
dien th se dan en s hnh thanh
isovaleryl-CoA. S chuyen hoa
tiep tuc cua isovaleryl- CoA hien
nh trat t c thc hien nh trat
t cac phan ng sau:
4 3 2 1
-CH
2
-CH
2
-CH-CO-S-CoA
CH
3
-oxy-hoa dan ti
s hnh thanh propionyl-CoA
4 3 2 1
-CH
2
-CH-CH
2
-CO-S-CoA
CH
3
-oxy-hoa b phong toa
4 3 2 1
CH
3
-CH-CH
2
-CO-S-CoA
CH
3
-oxy-hoa b phong toa va
phan con lai la isovaleryl-CoA
Hnh 10.1. Cac kieu phan nhanh cua
acid beo va quan he gia chung vi he
enzyme -oxy-hoa.

CH
3
-CH-CH
2
-CO-S-CoA CH
3
-C=CH-CO-S-CoA
CH
3
CH
3

FAD FAD.H
2

Biotin-COO
-
Biotin H
2
O
CH
3
-C=CH-CO-S-CoA

CH
2
-COO
-

OH Acetyl-CoA + Acid acetoacetic
CH
3
-C-CH
2
-CO-S-CoA NADP.H
CH
2
-COO
-
Acid mevalonic


Acid mevalonic la san pham trung gian cua nhieu qua trnh sinh tong hp, trong
o co qua trnh sinh tong hp cac hp chat steroid. S chuyen hoa tiep tuc cua acid
acetoacetic se c xem xet sau.
Propionyl-CoA hnh thanh trong qua trnh -oxy-hoa khong nhng cac acid beo
co mach carbon phan nhanh trong trng hp mo ta tren ay ma noi chung cac acid
beo co so le nguyen t carbon. Ngoai ra, goc propionyl 3 carbon cung xuat hien khi
phan huy cac isoprenoid, cac aminoacid isoleucine, threonine va methionine. Hn
na, con ngi con can s dung mot lng nho acid propionic von co mat trong mot so
thc pham. Propionate cung c cho vao banh my e ngan can s phat trien cua nam
moc. ac biet oi vi ong vat nhai lai propionate la mot nguon nang lng quan
trong do no c vi khuan phan giai cellulose trong da day tao ra cung vi acetate va
butyrate.

Khoa Sinh hoc

HNH 10.2. CAC CON NG D HOA PROPIONATE VA

c/ -Oxy-hoa acid beo khong no.
1/ Nh cac phan ng -oxy-hoa cua con ng (a) dan en s hnh thanh dan
xuat CoA cua semialdehyde malonic e tiep tuc b oxy hoa thanh malonyl-CoA va
cuoi cung b decarboxyl-hoa thanh acetyl-CoA.
2/ thc vat va nhieu vi sinh vat -Oxypropionyl-CoA von hnh thanh trong
cac phan ng -oxy-hoa khong chuyen hoa thanh dan xuat CoA cua semialdehyde
malonic ma theo con ng (b) b thuy phan thanh -oxypropionate e sau o b oxy-
hoa thanh semialdehyde malonic roi t o theo oan cuoi cung cua con ng (a)
chuyen hoa thanh acetyl-CoA.
3/ Mac du -oxy-hoa la mot con ng n gian, ong vat bac cao con ton
tai mot con ng (c) phc tap hn vi s tham gia cua vitamine B
12
. No c bat au
bang phan ng carboxyl-hoa propionate phu thuoc biotin va ATP. (S)-methylmalonyl-
CoA hnh thanh trong phan ng nay sau o chuyen thanh dang ong phan (R)-
methylmalonyl-CoA. Chat nay tiep tuc chuyen hoa thanh succinyl-CoA vi s tham
gia cua vitamine B
12
.
Tach CoA-SH ra khoi succinyl-
CoA se giai phong succinate t do,
ong thi tao ra mot phan t ATP e
bu lai phan t ATP a s dung.
Succinate bang con ng -oxy-hoa
bien thanh oxaloacetate roi b
decarboxyl-hoa thanh pyruvate. Cuoi
cung bang phan ng decarboxyl-hoa
pyruvate se chuyen hoa thanh acetyl-
CoA.
Acid beo khong no cung b
phan huy chu yeu bang con ng -
Oxy-hoa vi s tham gia cua hai
enzyme bo sung (hnh 10.3). Ta hay
lay acid linoleic lam v du. Sau 3
GS.TS. Mai Xuan Lng
H h 10 3 O h id b kh
vong -oxy hoa, cung vi 3 phan t
acetyl-CoA se xuat hien
3,4
-cis-
6,7
-
cis-enoyl-CoA. Mot isomerase ac
hieu
3
-cis-
3
-trans-enoyl-CoA
isomer-ase) chuyen dan xuat khong
no nay thanh -
2,3
-trans--
6,7
-cis-
enoyl-CoA, tao ieu kien e -oxy-
hoa tiep tuc c thc
hien. Cat tiep 2 phan t acetyl-CoA na se dan en tnh trang la lien ket oi th hai
vi cau hnh cis- chuyen ti v tr
2,3
. Enzyme enoyl-CoA hydratase cua -oxy-hoa
gan them phan t nc vao chat nay, lam xuat hien -oxyacyl-CoA vi cau hnh D.
Nh mot epimerase ac hieu no c chuyen thanh dang L e -oxy-hoa lai c
tiep tuc.
d/ - va - Oxy hoa
Trong cac mo ong vat qua trnh phan huy acid beo c thc hien chu yeu bang
con ng -oxy-hoa. Tuy nhien trong cac te bao thc vat, ac biet la trong hat
nay mam
oxy hoa acid beo thng c thc hien theo c che -oxy-hoa, tc bang cach
tach
lan lt cac oan 1C. Chi tiet cua
c che nay cha c sang to,
nhng ngi ta biet c rang giai
oan au cua no thng la hydroxyl-
hoa carbon - e tao ra D- hoac L-
oxyacid (hnh 10.4). L-2-Oxyacid
tiep tuc b oxy-hoa de dang co le
bang con ng dehydrogen-hoa e
tao ra -cetoacid va sau o
decarboxyl-hoa chat nay vi s tham
gia cua H
2
O
2
.
Hnh 10.4. Oxy hoa acid beo.
Hnh 10.4. - Oxy hoa acid beo
Trong khi o D-oxyaxid co xu hng tch
luy va thng co mat trong la
xanh. Tuy nhien, chung cung co the
b oxy-hoa nhng gi lai nguyen t
hydro co dau (*). ieu o cho thay co
mot con ng dehydrogen-hoa khac
von xay ra ong thi vi qua trnh
decarboxyl-hoa.
Hnh 10.4. -Oxy Hoa acid beo

-Oxy-hoa mc o nao o cung xay ra trong cac mo ong vat, ac biet la trong
nao. Trong nhng trng hp -oxy-hoa b phong toa do cac nhom -CH
3
trong mach
nhanh th -oxy-hoa se la mot con ng vong e khac phuc s phong toa nay.
Cac acid beo co chieu dai trung bnh, va trong mot so trng hp, cac acid beo
co chuoi carbon dai hn co the trai qua qua trnh -oxy-hoa e tao ra cac dan xuat
hydroxyacid; nhng dan xuat nay sau o thng chuyen hoa thanh cac acid -,-
dicarboxylic. Trat t cac phan ng nay c thc hien vi s xuc tac cua cac enzyme
trong vi the cua gan ma enzyme au tien la mot monooxygenase ma hoat ong cua no
can s tham gia cua NADP, O
2
va cytochrome P
450
. Sau khi hnh thanh, nhng acid
dicarboxylic nay co the b cat ngan t mot trong hai au cua chung bang c che -oxy-
hoa.
3. The cetone.
Khi acid beo co so chan nguyen t carbon b phan huy bang con ng -oxy-
hoa th san pham trung gian cuoi cung trc khi chuyen hoa hoan toan thanh acetyl-
CoA la hp chat 4C acetoacetyl-CoA:
O
CH
3
- C - CH
2
- C - S-CoA
O
Trong c the chat nay la mot san pham trung gian quan trong va co le ton tai
tren mot the can bang vi acetyl-CoA. No khong ch co the b phan giai thanh 2 phan
t acetyl-CoA e i vao chu trnh acid tricarboxylic ma con la mot chat tien than quan
trong trong qua trnh tong hp cac hp chat isoprenoid, trong o co cholesterol. Ngoai
ra, acid acetoacetic cung co y ngha rat quan trong v la mot trong cac thanh phan cua
mau. Khi can thiet, no se b decarboxyl-hoa thanh acetone nh acetoacetate
decarboxylase:
O O
CH
3
- C- CH
2
- COO
-
+ H
+
CH
3
- C - CH
3
+ CO
2

No cung co the b kh nh mot dehydrogenase phu thuoc NAD.H thanh acid L-
3-oxybutyric. Can lu y la cau hnh cua chat nay ngc vi cau hnh cua D-3-
oxybutyryl-CoA von hnh thanh trong qua trnh -oxy-hoa acid beo.
Tat ca 3 hp chat: acetoacetate, acetone va 3-oxybutyrate eu co ten chung la
the cetone. Nong o cua chung trong mau tang ang ke trong nhieu trng hp benh
ly khac nhau, ac biet la khi b benh tieu ng, cung nh khi nhn oi. Viec tang
nong o cac the cetone rat nguy hiem oi vi c the v no kem theo hien tng giai
phong ion H
+
, lam cho mau b acid hoa.
Mat khac, acetoacetate va oxybutyrate la nhng nguon nang lng quan trong
cho hoat ong cua c va cac mo khac trong nhng trng hp thieu glucid. e s
dung the cetone lam nguon nang lng, can phai chuyen hoa chung thanh acetyl-
CoA:


4. S dung lipid d tr cho muc ch sinh tong hp. Chu trnh
glyoxylate.
Glycerol va acid beo t do hnh thanh trong qua trnh thuy phan lipid d tr
trong hat co dau khi chung nay mam thng c dung e tong hp glucid va cac hp
chat khac. Qua trnh nay c thc hien thong qua chu trnh glyoxylate, bien dang cua
chu trnh acid tricarboxylic, sau khi glycerol va acid beo bang cac con ng rieng
cua mnh bien thanh acetyl-CoA (hnh 10.5).

S khac biet gia chu trnh glyoxylate va chu trnh acid tricarboxylic bao gom
cac phan ng c ong khung trong hnh 10.5. Isocitrate nh isocitratase phan giai
thanh succinate va glyoxylate. Glyoxylate sau o ket hp vi phan t acetyl-CoA th
hai vi s xuc tac cua malate synthase e tong hp nen malate. Trong khi o
succinate c a ra khoi chu trnh e c dung cho cac muc ch sinh tong hp.
H10.5. Chu trnh glyoxilate
ong vat bac cao khong ton tai chu trnh nay, trong khi o vai tro cua no
thc vat lai rat quan trong.
Cac enzyme cua chu trnh glyoxylate, ac biet la isocitratase va malate
synthase, nh v trong glyoxysome - mot bao quan ch c phat hien thc vat trong
nhng te bao co kha nang bien lipid thanh glucid. Tai ay khong co cac enzyme van
chuyen ien t va phan ln cac enzyme cua chu trnh acid tricarboxylic.


IX. SINH TONG HP ACID BEO.
1. Sinh tong hp acid beo no.
Acid beo no c tong hp trong te bao nh s xuc tac cua mot he thong
enzyme gan lien vi mot chat mang acyl ac hieu co ten la acyl carier protein,
thng c ky hieu la ACP, hay ACP-SH e lu y s ton tai cua nhom -SH trong
phan t cua no.
Nguyen lieu e tong hp acid beo la acetyl-CoA von hnh thanh trong ty the
do -oxy-hoa acid beo hoac do decarboxyl-hoa acid pyruvic hoac t mot so nguon
khac. Cac phan t acetyl-CoA c van chuyen ra te bao chat nh ket hp vi
oxaloacetate e tao ra citrate. Chat nay ra khoi ty the nh mot c che van chuyen ac
biet e roi lai b phan giai thanh acetyl-CoA va oxaloacetate nh mot enzyme phu
thuoc ATP.
Mac du nguyen lieu e tong hp acid beo la acetyl-CoA, song trc het no can
c carboxyl-hoa thanh malonyl-CoA nh acetyl-CoA carboxylase:
CH
3
-C-S-CoA + CO
2
+ ATP HOOC-CH
2
-C-S-CoA + AMP + PP
i

O O
Sau o qua trnh sinh tong hp acid beo c bat au bang trat t 6 phan ng
mo ta trong hnh 10.6.
Trong cac phan ng (1) va (2) acetyl-CoA va malonyl-CoA nh cac enzyme van
chuyen tng ng acetyl transacylase va malonyl transacylase chuyen hoa thanh
acetyl-ACP va malonyl-ACP. Hai chat nay trong phan ng (3) ket hp vi nhau nh
-cetoacyl-ACP synthetase thanh san pham 4C acetoacetyl-ACP, ong thi giai
phong CO
2
. Acetoacetyl-ACP di tac dung cua -cetoacyl-ACP reductase vi
coenzyme la NADP.H b kh trong phan ng (4) thanh D--oxybutyryl-ACP. (Can
lu y rang trong -oxy-hoa -oxybutyryl-CoA co cau hnh L). Trong phan ng (5) vi
s xuc tac cua enoyl-ACP hydratase -oxybutyryl-ACP b dehydrate-hoa thanh san
pham khong no crotonyl-ACP. Cuoi cung, trong phan ng (6) xuc tac bi crotonyl-
ACP reductase vi coenzyme la NADP.H crotonyl-ACP se b kh thanh butyryl-ACP.


Hnh 10.6. S o sinh tong hp acid beo
Butyryl-ACP lai ket hp vi malonyl-ACP theo c che cua phan ng (3) va qua
trnh c lap lai nh tren, lam cho phan t acid beo noi dai them mot oan 2 carbon
na, tao ra mot acyl-ACP cha 6 carbon (C
6
). Khi phan t acyl-ACP at c chieu
dai can thiet, ACP-SH se c tach khoi goc acid beo nh enzyme thuy phan
deacylase.
Trong phc he multienzyme tong hp acid beo cua te bao eucaryote ACP-SH
chiem v tr trung tam va tiep xuc trc tiep vi 6 enzyme con lai (hnh V.8).
Coenzyme cua ACP-SH bao gom phosphopantotenate va -mercaptoethylamin.
Thong qua goc phosphate cua phosphopantotenate, coenzyme nay lien ket ester vi
goc serine cua chuoi polypeptide ( E. coli chuoi polypeptide nay cha 77 goc
aminoacid va goc serine th 36 gan vi coenzyme). Trong khi o nhom -SH cua
mercaptoethylamine lien ket vi c chat va vi cac san pham trung gian trong qua
trnh sinh tong hp. Bang cach o cac coenzyme cua ACP-SH lam nhiem vu nh mot
canh tay e chuyen giao cac san pham trung gian t enzyme nay en enzyme khac.
Cac san pham trung gian nh lien ket kha chat che vi nhom -SH tan cung cua ACP
nen khong b chi phoi bi cac qua trnh trao oi chat khac trong bao tng ma tai o
qua trnh sinh tong hp acid beo c thc hien.
Trong te bao E. coli va vi khuan noi chung cac enzyme cua he thong
synthetase acid beo ch thc hien chc nang xuc tac mot cach rieng le.
Toc o sinh tong hp acid beo trong te bao co le c ieu hoa bi toc o hnh
thanh triacylglycerine va glycerophospholipid, v acid beo t do khong tch luy trong
te bao vi ham lng cao.
Ben canh c che tren ay con ton tai mot so c che khac, chu yeu e
noi dai them mach acid beo san co 12-16 nguyen t carbon: 1/ Trong ty the mach
carbon cua acyl-CoA c noi dai bang cach ket hp vi acetyl-CoA theo c che
ngc vi -oxy-hoa, tr phan ng hydrogen-hoa lien ket oi xay ra vi s tham gia

cua NADP.H ch khong phai cua FAD.H
2
. He enzyme nay cua ty the cung co the noi
dai mach carbon cua acid beo khong no. 2/ Trong vi the ton tai mot he thong enzyme
khac xuc tac qua trnh noi dai mach carbon cua acid beo no va khong no bang cach
gan them cac goc acetyl t malonyl-CoA. Phan ng hydrogen-hoa lien ket oi ay
cung c thc hien nh NADP.H.
e tong hp cac acid beo co so nguyen t carbon le, propionyl- ACP se thay
the cho acetyl-ACP tham gia phan ng (3) vi malonyl-ACP. Trong khi o, neu thay
cho acetyl-ACP la cac san pham d hoa cua valine, leucine va isoleucine, v du
isobutyryl-ACP, isovaleryl-ACP va - methylbutyryl-ACP, th mach acid beo c
tong hp se co cau truc phan nhanh tai au CH
3
tan cung, tng t nh cau truc phan
nhanh cua chnh nhng aminoacid nay. Trong trng hp methylmalonyl-ACP thay
cho malonyl-ACP trong phan ng (3) th cau truc phan nhanh dang cac nhom methyl
se co mat tai cac khu vc khac nhau cua mach acid beo.

Hnh 10.7. Phc he multienzyme synthetase acid beo trong te bao eucaryote
2. Sinh tong hp acid beo khong no.
cac sinh vat khac nhau co cac con ng khac nhau dan en s hnh thanh
lien ket oi trong acid beo khong no. Lien ket oi co the hnh thanh ngay trong qua
trnh tang trng mach carbon hoac sau khi qua trnh tang trng ket thuc (hnh 10.8).
E. coli va cac vi khuan tng t von song trong ieu kien k kh qua trnh
tong hp acid beo khong no c thc hien bang cach tao lien ket oi giai oan hnh
thanh -oxydecanoyl-ACP cha 10 nguyen t carbon. Di tac dung cua -

oxydecanoylthioester dehydratase nhom -hydroxyl c loai bo e tao lien ket oi
vi cau hnh -
3,4
-cis- (ch khong phai -
2,3
-trans- nh xay ra trong phan ng (5) cua
sinh tong hp acid beo no). Sau o nh he enzyme synthetase acid beo mach carbon
tiep tuc tang trng e tao ra palmitooeyl-ACP (C
16
) hoac varsenoyl-ACP (C
18
) rat
pho bien trong nhom te bao nay.
Ngi ta cho rang ong vat, thc vat va phan ln vi sinh vat hieu kh acid
beo khong no cha mot lien ket oi c hnh thanh t cac dan xuat coenzyme A cua
acid beo no nh mot loai oxygenase ac biet nh v trong vi the. V du:
Palmitoyl-CoA + NADP.H + H
+
+ O
2
Palmitooleyl-CoA + NADP
+
+ H
2
O

Tnh chat ac biet cua phan ng the hien cho phan t oxy c dung lam chat
nhan hai cap ien t: mot cap t c chat va mot cap t coenzyme. Nhng enzyme loai
nay c goi la "oxygenase co chc nang hon hp".
Hnh 10.8. Cac con ng sinh tong hp acid beo khong no.
Acid beo cha nhieu lien ket oi hnh thanh t stearoyl-CoA. Trong te bao
chat cua ong vat va thc vat t stearoyl-CoA tao ra oleoyl-CoA. Sau o t axit beo

khong no co mot lien ket oi nay vi s tham gia cua NADP.H, feredoxin va O
2
tao ra
linoleoyl-CoA va linolenoyl-CoA cung nh cac acid beo co nhieu lien ket oi hn.
ong vat oleoyl-CoA khong chuyen hoa c thanh linoleoyl-CoA, v vay cac acid
beo khong no cha nhieu lien ket oi nh acid linoleic,acid linolenic, acid arachidonic
khong c tong hp trong c the cua chung, do o nhng acid beo nay c goi la
"acid beo khong thay the", hay vitamin F.
X. SINH TONG HP TRIACYLGLYCERIN.
Triacylglycerin c tong hp t L-glycerol-3-phosphate va cac dan suat
coenzyme A cua acid beo.
Glycerol-3-phosphate c hnh thanh t hai nguon khac nhau:
1/ Kh dioxyacetonephosphate do glycerophosphate dehydrogenase xuc tac:
Dioxyacetonephosphate + NAD.H+H
+
Glycerol-3-phosphate + NAD
+

2/ phosphoryl-hoa glycerol nh glycerokinase:
Glycerol + ATP Glycerol-3-phosphate + ADP
Giai oan au cua qua trnh sinh tong hp triacylglycerin la phan ng tong hp
acid phosphatidic:

O

CH
2
OH O O H
2
-C-O-C-R
1

HO-C-H + R
1
-C-S-CoA + R
2
-C-S-CoA R
2
-C-O-C-H
CH
2
O-(P) O H
2
C-O-(P)
(Glycerophosphate) (Acid phosphatidic)
mot so vi sinh vat, v du E. coli, acid beo tham gia phan ng tren dang dan
xuat cua ACP-SH.
Nh mot phosphatase ac hieu acid phosphatidic loai bo goc phosphate e tr
thanh diacylglycerin. Cuoi cung, chat nay gan them goc acid beo th ba e tao ra
triacylglycerin.
Triacylglycerin c tong hp va tch luy trong cac mo d tr cua ong vat va
thc vat. Trong c the ong vat qua trnh tch luy nay c ay manh khi thc an co
ham lng glucid cao. Trong trng hp o ham lng glucose trong mau tang len va
ong thi tang mc o d tr glycogen trong te bao. Ham lng ATP cung tang len,
gay c che chu trnh acid tricarboxylic. Ket qua la acid citric b a ra khoi ty the va
b phan giai di tac dung cua enzyme citrate liase phu thuoc ATP:
Citrate + CoA-SH + ATP Oxaloacetate + Acetyl-CoA + ADP + P
i

Oxaloacetate co the b kh thanh malate va sau o b oxy-hoa va decarboxyl-
hoa thanh acid pyruvic roi xam nhap lai vao ty the; con acetyl-CoA c s dung e

tong hp acid beo va sau o cung vi glycerol-3- phosphate tong hp lipid d tr, lam
cho qua trnh tch luy m c tang cng.
Qua trnh tch luy m con c thuc ay do acid citric co tac dung hoat hoa
enzyme acetyl-CoA carboxylase. Insulin cung co tac dung thuc ay qua trnh tch luy
m do no lam tang hoat tnh cua he enzyme tong hp lipid, ac biet la enzyme citrate
liase phu thuoc ATP. Ngoai ra, thong qua tac dung c che s hnh thanh AMP vong
no ngan can qua trnh huy ong m d tr. Trong khi o cac hormone khac nh
glucagon, adrenalin kch thch s hnh thanh AMP vong va do o hoat hoa lipase, dan
en s tang cng huy ong m d tr.
XI. SINH TONG HP GLYCEROPHOSPHOLIPID VA
GLYCEROGALACTOLIDID.
vi khuan phan ln glycerophospholipid (hay phosphoglyceride, phosphatide)
c tong hp bang con ng chuyen hoa acid phosphatidic thanh CDP-
diacylglycerine. Sau o chat nay ket hp vi cac base khac nhau e tao ra
phosphoglyceride tng ng, ong thi giai phong CMP (hnh X.9). Phan ln cac phan
ng nay xay ra trong mang noi chat.

Hnh 10.9. S o cac phan ng sinh tong hp glycerophospholipid
ong vat phosphatidylcholine va phosphatidylethanolamine con co the c
tong hp bang con ng:
Choline + ATP Phosphorylcholine + ADP,
CTP + Phosphorylcholine cytidinediphosphatecholine + PP
vc

Cytidinediphosphatecholine + Diacylglycerine Phosphatidylcholine +
CMP.
Neu ethanolamine thay vao v tr cua choline trong cac phan ng tren th san
pham se la phosphatidylethanolamine.
Phosphatidylinoside thong qua cac nhom OH t do cua goc inositol co the nhan
t ATP mot, hai nhom phosphate va lien ke t vi no mot cach long leo, nh o
phosphatidyl-inoside thc hien c chc nang van chuyen phosphate qua mang.
Glycerogalactolipid cua luc lap c tong hp bang cach gan vao
diacylglycerine mot hoac hai goc galactose t UDP-galactose (hnh 10.10).

Hnh 10.10. S o cac phan ng sinh tong hp glycerogalactolipid
XII. SINH TONG HP SPHYINGOLIPID VA GLYCOLIPID.
Sphingolipid c tong hp t san pham ngng tu cua palmitoyl-CoA vi serine.
San pham ngng tu nay se b kh bi NADP.H thanh sphinganine
(dihydrosphingosine), sau o boac b oxy hoa bi flavoprotein thanh sphingosine hoac
hydroxyl hoa thanh phytosphingosine (hnh 10.11).

Hnh 10.11. S o cac phan ng sinh tong hp sphigolipid.

Sphingosine bang nhom NH
2
cua mnh se ket hp vi mot acyl-CoA e tao ra
seramide. Chat nay tac dung vi CDP-choline se lam xuat hien sphingomieline hoac
vi cac dan xuat UDP va CDP cua cac saccharide khac nhau se tong hp nen
glycolipid (serebroside va ganglioside) (hnh 10.12).

XIII. SINH TONG HP STERINE.
Trong c the ong vat, thc vat va nam sterine c tong hp t acetyl-CoA. (
vi khuan khong co nhom hp chat nay).
Cholesterine ong vat la tien than cua acid mat va cua cac loai hormone
steroid. Qua trnh sinh tong hp cholesterine kha phc tap. No bao gom nhng bc
chu yeu sau ay:
1/ Ngng tu hai phan t acetyl-CoA e tao nen acetoacetyl-CoA;
2/ Acetoacetyl-CoA tac dung vi phan t acetyl-CoA th ba e tao nen -oxy--
metylglutaryl-CoA;
3/ Mot trong hai nhom carboxyl cua -oxy--metylglutaryl-CoA b kh e tao ra
acid mevalonic:
OH
HOOC CH
2
C CH
2
CO ~ S CoA + 2NADP.H + 2H
+

CH
3

OH
HOOC CH
2
C CH
2
CH
2
OH + 2NADP
+
+ CoA-SH
CH
3

4/ Acid mevalinic c phosphoryl hoa thanh mevalonylpyrophosphate:
OH OH
HOOCCH
2
CCH
2
CH
2
OH +2ATPHOOCCH
2
CCH
2
CH
2

O- P -O-P +
2ADP
CH
3
CH
3
5/ Decarboxyl hoa mevalonylpyrophosphate thanh isopentenylpyrophosphate:
Mevalonylpyrophosphate +ATP CH
2
=CCH
2
CH
2

O P O - P +

Hnh 10.12. S o cac phan ng sinh tong hp
Sphingomieline , serebroside va ganglioside

CH
3
+CO
2
+H
2
O+ADP+P
vc

6/ Ngng tu 3 phan t isopentenylpyrophosphate e tao ra
farnesylpyrophosphate:
CH
3
C=CHCH
2
CH
2
C=CHCH
2
CH
2
C=CHCH
2
CH
2

O P O P
CH
3
CH
3
CH
3
7/ Phan t farnesylpyrophosphate ngng tu vi dang ong phan cua no la
nepolidolpyrophosphsate e tao nen squalen:
8/ Chuoi hydrocarbon cua squalene chuyen hoa thanh dang mach vong, gan
them nhom OH C
3
va bien thanh lanosterine;
9/ Sau mot so khau trung gian lanosterine bien thanh cholesterine;
Hai khau sau cung xay ra tren he thong mang cua mang noi chat. Trong khi o
cac phan ng thuoc cac giai oan trc xay ra trong bao tng.
C che sinh tong hp sterine thc vat va nam men (stigmasterine, ergosterine
v.v...) ve c ban cung giong vi c che tong hp cholesterine.


CHNG 6. NUCLEOTIDE VA ACID NUCLEIC

Cung vi protein, acid nucleic ong vai tro cc ky quan trong trong viec bao ton
s song. Tuy nhom hp chat nay a c phat hien hn 100 nam ve trc, cau tao,
tnh chat va chc nang cua chung ch mi c hieu biet mot cach sau sac t nhng
nam 50 cua the ky 20 nh ng dung nhng phng phap nghien cu vat ly va hoa hoc
chnh xac.
Tren c s s khac biet ve thanh phan hoa hoc, acid nucleic c chia thanh hai
nhom ln. o la acid ribonucleic (ARN) va acid deoxyribonucleic (ADN). ADN co
trong lng phan t t vai trieu en vai tram trieu, la thanh phan khong the thieu c
cua nhiem sac the trong nhan te bao. Ngoai ra, ADN con co mat trong ti the va luc
lap. ARN co nhieu loai: ARN van chuyen (tARN) co trong lng phan t tng oi
nho (25.000 35.000); ARN ribosome (rARN), noi chung, co trong lng phan t kha
ln (t 1,7 en 1,2 trieu), tr mot vai loai ch ln hn tARN mot t; ARN thong tin
(mARN) co trong lng phan t t 300.000 en 4 trieu; ARN virus co trong lng
phan t t 1 en 2 trieu. Ngoai ra, con co mot so ARN khac, chu yeu la ARN phan t
nho ma cau truc va chc nang cua chung cha c hieu biet nhieu.
ai bo phan acid nucleic (ca ADN va ARN) la nhng biopolymer dang si hnh
thanh t cac n v cau tao (monomer) co ten chung la (mono)nucleotide. Moi
nucleotide c cau tao t ba thanh phan: monosaccharide, base nit va acid
phosphoric.
I. NUCLEOTIDE.
Phan ln nucleotide, hay con goi la mononucleotide, la n v cau tao cua acid
nucleic. Chung oc cau tao t base nit, ng pentose va acid phosphoric.
Tat ca cac nucleotide tham gia cau
tao nen acid nucleic eu cha mot trong
hai loai monosaccharide: -D-ribose hoac
2--D-deoxyribose.
Ca hai loai pentose nay eu co
dang cau truc vong furanose. Can lu y rang, khi dang t do cac nguyen t carbon
trong phan t c anh so t 1 en 5, nhng khi tr thanh mot bo phan cua phan t
nucleotide, chung phai c anh so t 1' en 5' (e phan biet vi cac nguyen t
carbon trong base nit).
Ngoai ribose va deoxyribose, ribitol - san pham kh cua ribose - cung co the
tham gia trong thanh phan cau tao cua mot so nucleotide ac biet.
Trong hai loai acid nucleic - acid ribonucleic va acid deoxyribonucleic - co 5
loai base nit thng gap, 3 loai chung cho ca ADN va ARN (adenine, guanine va

cytosine), loai th t (uracil) ac trng cho ARN, con loai th 5 (thymine) hau nh ch
co mat trong ADN.

Hnh IV.1. Cong thc cau tao cua cac base nit chu yeu.

Adenine va guanine la dan xuat cua purine va do o c xep vao nhom base
purine; 3 base con lai la dan xuat cua pyrimidine va do o c xep vao nhom base
pyrimidine.
Thymine cung c xem
la mot base th yeu
trong thanh phan cau tao
caa ARN; ngc lai,
uracil la base th yeu
trong ADN. Cau truc cua
cac base chu yeu c
trnh bay trong hnh
IV.1. Cach anh so cac
nguyen t carbon va nit
trong moi phan t base
c gii thieu qua cac
ai dien la purine va
pyrimidine. T cong
thc cau tao cua cac
base chu yeu ta co the
de dang viet cong thc
Hnh IV.2. Hien tng ho bien cac base cha oxy

cau tao cua cac base th
yeu tren c s ten goi
cua chung.

ac iem quan trong cua cac dan xuat cha oxy cua purine va pyrimidine la kha
nang ho bien, tc chuyen hoa tng ho gia cac dang enol (lactim) va cetone (lactam)
(hnh IV.2):
Ngoai cac base chu yeu noi tren, trong mot so loai acid nucleic, ac biet la
trong ARN van chuyen (tARN) va ARN ribosom (rARN), con hay gap mot so base
khac vi ham lng khong ln. Chung c goi la cac base th yeu, phan ln la cac
dan xuat hydrogen hoa, methyl hoa, oxymethyl hoa cua cac base chu yeu, v du
dihydrouracil (UH
2
),2-methyladenine (A-CH
3
), 5-oxymethyl-cytosine (C-OCH
3
) v.v.
Tat ca cac base nit eu hap thu anh sang manh nhat trong vung pho cc tm.
Tnh chat nay c ng dung e xac nh ham lng acid nucleic va cac san pham
thuy phan cua chung.
Khi mot base nit lien ket vi mot pentose bang lien ket N-glycoside se tao ra
cac san pham co ten chung la nucleoside. Tuy thuoc cho loai pentose nao va loai
base nao tham gia cau tao nen nucleoside ma moi loai co ten goi rieng cua mnh. Tren
c s thanh phan pentose ngi ta phan biet hai loai nucleoside la ribonucleoside
(pentose la ribose) va deoxyribonucleoside (pentose la deoxyribose). cach nhn
khac, cac nucleoside lai c phan biet la purine nucleoside hay pyrimidine nucleoside
tuy thuoc cho base purine hay base pyrimidine tham gia trong thanh phan cau tao
cua chung. Can lu y rang trong cac purine-nucleoside lien ket glycoside hnh thanh
gia C-1' vi N-9, trong khi o cac pyrimidine nucleoside - gia C-1' vi N-1. Cong
thc cau tao cua mot so nucleoside ien hnh c gii thieu trong hnh VI.3. Thong
qua cac cong thc cau tao nay ta co the hnh dung cong thc cau tao cua cac
nucleoside khac.
Nucleoside c goi ten theo base nit theo nguyen tac:
- Neu base la dan xuat cua purine th uoi "- ine" c oi thanh "-osine", v du:
adenosine, guanosine;
- Neu base la dan xuat cua pyrimidine th uoi "-acil" hoac "-ine" c oi thanh
"-idine", v du: cytidine, uridine, thymidine.
Pseudouridine () trong hnh IV.3 la mot trng hp ac biet, hau nh ch gap
trong tARN vi t le rat thap.

T cac cong thc cau tao trong hnh VI.3 ta co the hnh dung cong thc cau tao
cua cac nucleotide khac vi s tham gia cua guanine, cytidine, uracil, thymin v.v...

Hnh IV.3 . Cau tao cua mot so nucleoside ien hnh.
Khi mot nucleoside ket hp vi
mot goc phosphate bang lien ket
ester thong qua mot trong cac nhom -
OH con lai cua goc pentose se tao ra mot
nucleotide tng ng. ieu o co ngha la
moi ribonucleoside co the tao ra ba loai
nucleotide, trong o goc phosphate co the
gan tai C-2', C-3' hoac C-5'; trong khi o
moi deoxyribonucleoside ch tao ra hai
loai nucleotide tng ng v tai C-2' khong
co nhom -OH e tng tac vi phosphate.
Tuy thuoc vao v tr gan goc phosphate ma
san pham c goi la 2'-, 3'- hay 5'-
nucleotide. Chung cung con c goi
tng ng la2'-, 3'- hay 5'-nucleoside
monophosphate, viet tat la 2'-NMP,
3'-NMP va 5'-NMP. Trong te bao chu
yeu ton tai cac loai 3'- va 5'-
Hnh IV.4. Cau tao cua cac loai 5'-
, 3'-va 2'-nucleotide

nucleotide. Cau tao cua cac loai
nucleotid nay c gii thieu trong
hnh IV.4 vi base nit la adenine.
Trong bang IV.1 gii thieu ten goi va cach viet tat cua mot so ribonucleoside va
ribonucleotide pho bien. Trong trng hp goc mono-saccharide la deoxyribose, ten
goi cua cac nucleoside va nucleotide c them tiep au ng "deoxy-", va trc cac
ky hieu viet tat them ch "d", v du deoxyadenosine, acid deoxyadenylic,
deoxyadenosine monophosphate, dAMP, dA.
Bang IV.1. Ten goi va cach viet tat cua mot so nucleotide.
B
as
e
ni
t
Nucle
oside
Nucleotide Viet tat
Adenine Adenosine
Acid adenylic
Adenosine monophosphate
AMP, A
Guanine Guanosine
Acid guanilic
Guanosine monophosphate
GMP, G
Cytosine Cytidine
Acid cytidilic
Cytidine monophosphate
CMP, C
Uracil Uridine
Acid uridilic
Uridine monophosphate
UMP, U
Thymine Thymidine
Acid thymidilic
(Deoxy)Thymidine momophosphate
dTMP, T
Ngoai cac nucleotide gii thieu tren ay co chc nang chu yeu la tham gia cau
tao nen cac ai phan t acid nucleic, con co mot so nucleotide khac co cac vai tro
quan trong khac trong i song cua te bao (hnh IV.5).
Truoc IeI, do Ia cac nucIeoIIde vong. NIung nucIeoIIde IoaI nay InI
IIanI kII goc pIospIaIe IIen keI esIer dong IIoI voI IaI nIom -OH cua goc
rIbose. V du dIen InI Ia IaI IoaI AMP vong (cAMP). o Ia 2'-'-cAMP va '-
5'-AMP. '-5'-cAMP dong vaI Iro quan Irong Irong moI so qua IrnI dIeu Ioa
Irao doI cIaI, con 2'-'-cAMP Ia san pIam Irung gIan cua qua IrnI pIan
gIaI ARN duoI Iac dung cua moI so enzyme rIbonucIease.
Nhom nucleotide ac biet th hai la cac nucleoside polyphosphate, bao gom
nucleoside diphosphate (NDP), v du ADP, va nucleoside triphosphate (NTP), v du
ATP. Tnh chat ac biet cua cac NDP va NTP la cho mot hoac hai goc phosphate
na c gan vao phan t nucleoside monophosphate bang cac lien ket giau nang
lng (lien ket cao nang) ma ngi ta thng ky hieu bang dau ~, nh mo ta trong
hnh IV.5. Nh s ton tai cua cac lien ket cao nang nay nen cac NDP va ac biet la

cac NTP ong vai tro quan trong trong trao oi nang lng cua te bao va tham gia
hoat hoa nhieu hp chat trung gian cua cac qua trnh trao oi chat.
Nhom nucleotide ac biet th ba bao gom nhng hp chat ma thanh phan base
nit va monosacchride cua chung thng khong giong nh a mo ta tren. V du ien
hnh cho nhom nucleotide ac biet nay la nicotinamide mononucleotide (NMN),
flavine mononucleotide (FMN) va coenzyme A (CoA-SH) ma cong thc cau tao cua
chung c gii thieu trong hnh IV. 5. Do co kha nang oxy hoa-kh thuan nghch,
nen NMN va FAD tham gia trong hang loat cac enzyme oxyhoa-kh vi t cach la
coenzyme. Trong khi o CoA-SH ong vai tro rat quan trong trong trao oi lipid va
mot so qua trnh trao oi chat khac. NMN va FMN con la thanh phan cau tao cua cac
coenzyme oxyhoa-kh phc tap hn. o la cac dinucleotide NAD
+
, NADP
+
va FAD
(hnh IV.6).



Hnh IV.5. Cau tao cua mot so nucleotide co chc nang ac biet

Hnh IV.6. Cau tao cua NAD
+
, NADP
+
va FAD.
II. POLYNUCLEOTIDE
C s cau truc cua acid nucleic la cac chuoi polynucleotide cau tao t nhieu n
v (mono)nucleotide. Trong nhng chuoi nay cac mononucleotide noi vi nhau bang
cac lien ket 3'-5'-phosphodiester nh mo ta trong hnh 4.7. Cac chuoi polynucleotide
thng cha t hang chuc en hang tram goc mononucleotide. Tuy nhien, cung co
nhng chuoi polynucleotide ngan, cha khong qua 10 goc nucleotide va chung c
goi chung la oligonucleotide (bao gom di-, tri-, tetra-, penta-,hexanucleotide v.v...).

Ngi ta phan biet hai loai polynucleotide la polyribonucleotide - c s cau truc
cua ARN, va polydeoxy-ribonucleotide - c s cau truc cua ADN. Trong hnh IV.7A
gii thieu mot oan polynucleotide e ta co the hnh dung s hnh thanh lien ket 3'-5'-
phosphodiester. e mo ta thanh phan va trat t sap xep cua cac goc nucleotide
trong nhng chuoi polynucleotide ngan, ngi ta thng dung kieu mo hnh n gian
nh trnh bay trong hnh IV.7B. Kieu mo ta nay cho phep phan biet hai thanh phan
trong lien ket phosphodiester, tc lien ket a noi goc phosphate vi C-3' va lien ket b
noi goc phosphate vi C-5', qua o tao ra cau noi 3'-5'-phosphodiester gia hai
nucleotide ke can. Viec phan biet lien ket a vi lien ket b co y ngha rat quan trong do
hai kieu lien ket nay mang tnh ac hieu khac nhau oi vi cac enzyme thuy phan acid
nucleic. oi vi cac chuoi poly-nucleotide dai, khong the s dung kieu mo ta nay v se
rat cong kenh, va ngi ta thng dung kieu mo ta bang day ch cai A, G, C, U hoac
T, tc ch cai au tien cua ten tieng Anh cac base nit nh trnh bay qua v du trong
hnh IV.7C. Trong trng hp oi vi ADN trc day ch cai nay, tc au tan cung
5 cua chuoi polypeptide, viet them ch d (deoxy-), v du dpApGpTp...

Hnh IV.7. Cau tao cua polynucleotide

III. ADN, NHIE M SAC THE VA MAT MA DI TRUYEN.
Cac chuoi polydeoxyribonucleotide cua ADN c cau tao t 4 loai nucleotide
la dAMP, dGMP, dCMP va dTMP. Ngoai ra, oi khi ngi ta con tm thay mot lng
nho cac dan xuat methyl-hoa cua cac nucleotide nay, v du 6-methytladenine, 5-

methylcyto-sine v.v... c tm thay trong nhieu loai ADN cua vi khuan, ong vat va
thc vat. Thanh phan va trat t sap xep cua nucleotide trong cac chuoi
polydeoxyribonucleotide, tc cau truc bac 1 cua ADN vo cung a dang. S a dang
nay chnh la c s cua tnh a dang cua the gii sinh vat, bi v, nh ta se thay sau
nay, no la c s cua qua trnh tien hoa va lien quan mat thiet vi tnh di truyen.
e hieu ro chc nang sinh hoc cua ADN, ben canh nhu cau xac nh cau
truc bac 1 con can phai hieu ro cau truc khong gian cua chung. Mo hnh cau truc
khong gian cua phan t ADN c xay dng tren c s hang loat nghien cu trong
lnh vc sinh hoc phan t, trong o quan trong nhat la cac cong trnh cua Chargaff va
cua Franklin va Wilkins.
Sau nhieu nam nghien cu (1949-1953), Chargaff va nhng ngi cong tac cua
ong a neu len nhng ket luan quan trong ve ac iem hoa hoc cua ADN trong te bao.
Nhng ket luan nay ngay nay c moi ngi cong nhan va goi la cac quy luat
Chargaff. o la:
1/ Cac che pham ADN tach t cac mo khac nhau cua cung mot c the co thanh
phan nucleotide nh nhau;
2/ Thanh phan nucleotide cua ADN trong c the thuoc cac loai khac nhau la
khong giong nhau;
3/ Thanh phan nucleotide cua ADN trong c the thuoc mot loai nao o khong
phu thuoc vao tuoi, ieu kien dinh dng va ieu kien moi trng;
4/ Hau nh trong tat ca cac che pham ADN a nghien cu so goc adenine bang
so goc thymine (A = T), con so goc guanine bang so goc cytosine (G = C). ieu o
dan en so goc purine bang so goc pyrimidine (A+G = C+T);
5/ ADN cua cac loai von co quan he ve mat he thong hoc cang gan nhau th co
thanh phan nucleotide cang giong nhau; con cac loai cach xa nhau trong qua trnh tien
hoa th khac nhau kha ro ve thanh phan nucleotide. Co ngha la thanh phan nucleotide
cua ADN co the c s dung nh mot trong nhng c s cua phan loai hoc.
Cac nha khoa hoc nay cung phat hien c rang ong vat va thc vat, tc
nhng c the bac cao ch co ADN thuoc kieu AT, tc A+T > G+C, trong khi o ADN
vi khuan gom ca hai kieu AT va GC.
Ham lng nh nhau (tnh theo mol) cua mot so base trong ADN cho phep gia
nh rang net cau truc ac trng cua ADN la s ton tai nhng moi quan he hoan toan
xac nh gia so lng cac base khac nhau.
Song song vi phat hien cua Chargaff, ket qua phan tch cau truc bang tia Rn-
ghen do Franklin va Wilkins thc hien trong nhng nam 1950-1953 vi nhng che

pham ADN tinh khiet cho thay ADN co the ton tai hai dang A va B vi mc o
hydrat hoa khac nhau.
Tren c s cac ket qua nghien cu cua Chargaff cung nh cua Franklin va
Wilkins nam 1953 Watson va Crick a e xuat mot mo hnh cau truc khong gian cua
phan t ADN. Theo mo hnh nay dang cau truc B cua phan t ADN c cau tao bi
hai chuoi polydeoxy-ribonucleotide xoan phai, song song va ngc chieu nhau, nam
song oi xung quanh mot truc chung, tao thanh mot si xoan kep. Cac nguyen t
phosphore nam cach truc 1,0nm. Tren moi chuoi cac base mam cach nhau 0,34nm.
Theo tnh toan ban au mot vong xoan tron ven cha 10 cap base va co chieu dai
chieu len truc bang 3,4nm. Tuy
nhien cac phep o sau o cho thay mot vong xoan hoan chnh cha 10,5 cap base va
do o co chieu dai chieu len truc bang 3,6nm. (hnh IV.8). Dang B c xem la dang
on nh nhat trong ieu kien sinh ly. No ac trng cho trang thai co mc o hydrate
hoa cao.
Neu giam mc o hydrate hoa hoac nam trong cac moi trng tng oi co tnh
k nc, ADN se chuyen sang dang cau truc A, trong o cac base cach nhau 0,23nm
va moi vong xoan cha 11 cap base. Truc trung tam cua phan t ADN xoan kep khong
con thang goc vi cac mat phang cua cac cap base ma tao ra vi chung mot goc 20
o
,
ong thi cung khong xuyen qua cac mat phang nay ma ma nam lech sang pha nay
hoac pha khac. ac iem cau truc nay lam cho dang A ngan hn va co ng knh
ln hn dang B.
Cac hoa chat dung e ket tinh ADN thng lam cho no b dehydrate hoa, lam
cho ADN co xu hng ket tinh dang cau truc A.
Phan t ADN xoan kep tham ch co the ton tai dang quay trai (dang Z).
dang nay cac cap base cach nhau 0,38nm va 12 cap base tao nen mot vong
xoan hoan chnh. Bo khung cua phan t co dang zig-zak. Mot so trat t nucleotide de
tao ra dang cau truc Z hn la nhng trat t khac. Ngi ta cha biet ro vai tro sinh hoc
cua dang cau truc nay, ch biet rang cau truc Z c hnh thanh khi cac base cytosine
c methyl hoa. Methyl hoa la hien tng sinh hoc rat pho bien oi vi ADN, oi
vi eukaryote no co vai tro quan trong trong viec ieu hoa hoat tnh cua gen. Trong te
bao eukaryote co the ay la mot c che kiem tra hoat ong cua gene hoac tham gia
trong qua trnh cai bien gene (genetic recombination).



Hnh IV.8. S o mo ta s khac nhau gia cac dang cau truc A, B va Z cua
phan t ADN xoan kep

HNH IV.9. LIEN KET HYDRO GIA CAC CAP BASE BO
SUNG A T VA G C
Chuoi xoan kep c hnh thanh nh tnh chat bo sung gia cac base A vi T va
G vi C. Trong toan bo phan t moi base nit cua choi nay noi vi base bo sung
chuoi kia thong qua cac lien ket hydro (hnh IV-9).

Do tnh chat bo sung nay gia cac base nit ma trong phan t ADN xoan kep si
n nay co cau truc hoan toan bo sung vi si n kia.
Trong mach xoan kep cua ADN cac base nit do co tnh k nc nen nam ben
trong va khong tiep xuc vi nc, con cac nhom phosphate cung nh cac goc pentose
nam mat ngoai va tiep xuc trc tiep vi nc. Nh vay, cau truc xoan kep cua ADN
c on nh khong nhng nh lien ket hydro gia cac base bo sung ma con nh lien
ket k nc gia cac base nit vi nhau doc theo toan bo chieu dai phan t (tng tac
Steking).
Mo hnh cua Watson va Crick cho phep giai thch c che tai tao thong tin di
truyen mot cach chnh xac. Nh tnh chat bo sung cua hai mach n ma trat t
nucleotide cua mach nay xac nh trat t nucleotide cua mach kia. Song song vi viec
neu len mo hnh nay cac tac gia con gia thuyet rang phan t ADN c nhan oi bang
cach hai mach n cua phan t xoan kep tach ra va moi mach lam khuon e uc nen
mach mi bo sung vi no. Ket qua la hnh thanh hai phan t ADN giong het nhau,
trong moi phan t cha mot mach me va mot mach con. C che nhan oi nay cua
ADN c goi la c che ban bao thu.
Ben canh cau truc si xoan kep mo ta tren ay, mot so loai sinh vat ac biet
con co cac dang cau truc khac cua ADN. V du, E. coli va nhieu vi khuan khac co
dang cau truc xoan kep mach vong do hai au cua moi mach n noi vi nhau bang
lien ket phosphodiester. Dang cau truc nay cung c ghi nhan trong ti the, luc lap
cua te bao eukaryote va nhieu loai virus, v du bacteriophag ; trong khi o
bacteriophag X 170 ADN lai co dang cau truc vong si n.
Tat ca cac phan t ADN trong phan ln thi gian ton tai trong te bao trang thai
sieu xoan do tac dung cua cac enzyme ac biet. Trong khi o mot so enzyme khac lai
lam giam mc o sieu xoan, tao ieu kien e phan t ADN nhan oi hoac sao chep
ma di truyen sang cac phan t ARN. Nang lng tch luy trong trang thai sieu xoan
cua phan t ADN khi can thiet se c s dung cho qua trnh thao xoan cuc bo e
phan t ADN thc hien chc nang cua mnh.
Phan t ADN xoan kep di tac dung cua nhiet o cao, cua pH thai cc cua moi
trng, cua hang so ien moi thap va cua cac yeu to pha v lien ket hydro khac nh
urea, amide cua cac acid carboxylic... co the tach ri thanh hai si n rieng biet vi
ket cau hon on. Hien tng nay c goi la s bien tnh cua ADN. Neu dan dan loai
tr tac nhan gay bien tnh trong khi phan t ADN cha bien tnh hoan toan (mot phan
phan t van con ton tai trang thai xoan kep), cau truc xoan kep nguyen thuy cung
vi ay u cac tnh chat hoa hoc va sinh hoc cua no co the c khoi phuc hoan toan.
Trong trng hp nay ta co bien tnh thuan nghch. Neu phan t ADN a b bien tnh
hoan toan va loai tr tac nhan gay bien tnh mot cach ot ngot, cau truc xoan kep
nguyen thuy cua no se khong khoi phuc c na, tc phan t ADN a b bien tnh
khong thuan nghch.

S bien tnh cua ADN kem theo nhng thay oi ang ke cac tnh chat vat ly cua
no, ac biet, o hap thu tia t ngoai ( = 260nm) tang len. Hien tng nay c goi la
hieu ng u sac. Nguyen nhan cua no la do tnh nhc sac cua ADN xoan kep, tc o
hap thu tia t ngoai cua ADN nguyen thuy (xoan kep) nho hn tong o hap thu cua
cac base purine va pyrimidine khi chung ton tai trang thai t do. Khi b bien tnh, o
hap thu tia t ngoai cua ADN si n tang len 20 60%, v mc hap thu cua chung
bang tong o hap thu cua so lng tng ng cac base nit t do. Hieu ng u sac cua
ADN xoan kep lien quan trc tiep en cap base A-T. Ham lng cap base nay trong
ADN cang cao, hieu ng u sac cang ln. Nh vay, tren nguyen tac, co the xac nh
thanh phan nucleotide cua ADN bang cach xac nh hieu ng u sac cua no khi b
bien tnh.
Khac vi protein, ADN b bien tnh trong mot pham vi nhiet o rat hep. S bien
oi ot ngot nay tng t nh s nong chay cua cac tinh the hu c. V vay s bien
tnh v nhiet cua ADN thng c goi la s nong chay.
Hien tng bien tnh thuan nghch cua ADN c cac nha khoa hoc rat chu y,
bi v khong loai tr kha nang hien tng nay ong vai tro nhat nh trong cac qua
trnh nhan oi ADN va sinh tong hp ARN.
Trong qua trnh chiet rut va tinh che phan t ADN rat de b t, v vay viec thu
nhan ADN dang nguyen thuy va xac nh trong lng phan t cua no gap rat nhieu
kho khan. Tuy nhien, ngay nay cong viec nay a va ang thu nhan c nhng thanh
tu ang ke. V du, ngi ta a xac nh c phan t ADN duy nhat cua E. coli dai
1200 m, cha 4,2 trieu cap nucleotide va co trong lng phan t 2800 trieu. Si ADN
nay co cau truc vong, tao nen nhiem sac the duy nhat cua te bao vi khuan.
Nhiem sac the cua nhan te bao eukaryote c cau tao bi mot so si ADN
khong lo, trong nhieu trng hp co the dai en 2 met vi khoang 5,5 t cap
nucleotide do phai cha mot lng thong tin rat ln. Mot trong nhng thanh tu vo
cung v ai cua hoa sinh hoc va sinh hoc phan t cua the ky 20 la a xac nh c
cau truc bac I cua hau het ADN trong c the con ngi.
Nhiem sac the te bao eukaryote c cau tao t mot loai nucleoprotein co ten la
chromatine. Thanh phan chu yeu cua chromatine bao gom ADN, histone va mot so
protein khong phai histone. Histon la mot nho m protein co tnh base do cha nhieu
cac aminoacid co tnh base la lysine va arginine. Da tren t le gia Lys va Arg ngi
ta chia histone thanh 5 nhom. Giau lysine nhat la histone H
1
, giau arginine nhat la
histone H
4
; histone thuoc cac nhom H
2
A, H
2
B va H
3
lan lt chiem cac v tr trung
gian vi t le Lys/Arg giam dan.Thanh phan protein khong phai histone kha a dang
ve tnh chat va chc nang.
Chromatine co cau truc rat a dang. No bao gom cac cau truc hat goi la
nucleosome noi vi nhau bi cac oan nucleoprotein ngan. oan nucleoprotein nay
cau tao bi mot oan ADN dai khoang 200 cap nucleotide ket hp vi mot phan t

histone H
1
. Hat nucleosome c hnh thanh t oan ADN dai khoang 140-150 cap
nucleotide bao boc xung quanh phan loi vi 4 loai histone nhom H
2
A, H
2
B, H
3
va H
4
,
moi loai 2 phan t (hnh IV.10).

Hnh IV.10. Cau tao cua nucleosome
Ngay nay ngi ta a xac nh c rang thong tin di truyen c ma hoa trong
cac bo ba nucleotide cua ADN. Nhng bo ba nay c goi la bo ba mat ma hay codon,
lam nhiem vu ieu khien trat t aminoacid trong qua trnh sinh tong hp protein. V
moi codon gom ba trong bon loai nucleotide nen tong so codon bang 4
3
= 64. Ngay
nay y ngha cua tat ca 64 codon nay a c xac nh (bang IV.2). Y ngha nay ung
cho moi c the moi bac thang tien hoa.
Bang IV.2 . Mat ma di truyen.
Ch cai
th nhat
(au 5)
Ch cai th hai
U C A G
Ch cai
th ba
(au 3)

U

Phe
Phe
Leu
Leu
Ser
Ser
Ser
Ser
Tyr
Tyr
Term
Term
Cys
Cys
Term
Trp
U
C
A
G

C

Leu
Leu
Leu
Leu
Pro
Pro
Pro
Pro
His
His
Gln
Gln
Arg
Arg
Arg
Arg
U
C
A
G

A

Ile
Ile
Ile
Met
Thr
Thr
Thr
Thr
Asn
Asn
Lys
Lys
Ser
Ser
Arg
Arg
U
C
A
G

G

Val
Val
Val
Val
Ala
Ala
Ala
Ala
Asp
Asp
Glu
Glu
Gly
Gly
Gly
Gly
U
C
A
G


IV. ARN.
Khac vi ADN, phan t ARN thuoc tat ca cac loai c hnh thanh t mot chuoi
polynucleotide duy nhat. Tuy nhien, trong nhng ieu kien nhat nh tai tng khu vc
rieng biet cua chuoi polynucleotide nay co the hnh thanh cau truc xoan kep tren c s
tnh bo sung A-U va G-C.
Ham lng ARN trong a so te bao nhieu hn ADN gap nhieu lan. Trong te bao
vi khuan ARN ton tai trong te bao chat; trong te bao eukaryote ARN co mat trong
nhan, ty the, luc lap, ribosome va bao tng.
Trong te bao prokaryote cung nh eukaryote co ba nhom ARN chnh la ARN
thong tin (mARN), ARN van chuyen (tARN) va ARN ribosome (rARN). Ngoai ra, con
mot so loai ARN khac ma chc nang sinh hoc cua chung cha c xac nh. Mot
nhom ARN ac biet khac con co mat trong nhieu loai virus, ac biet la virus thc vat
va bacteriophage, o chung ong vai tro la vat chat di truyen (thay cho ADN).
1.ARN thong tin (mARN).
Phan t mARN ch cha 4 nucleotide la A, U, G va C.
mARN Irong Ie bao cI cIIem kIoang vaI pIan Iram ARN Iong so,
song bao gom Iang ngan IoaI kIac nIau voI Irong Iuong pIan Iu dao dong
Iu vaI Iram ngan den Iang IrIeu. PIan Iu cua cIung Iau nIu cI cIua cac
base A, O, C va U, raI I kII co maI cac base IIu yeu.

Cung nIu IaI ca cac IoaI ARN
kIac, mARN duoc Iong Iop bang
cacI sao cIep Iu cac gen Iuong ung
Iren cac pIan Iu ADN Irong qua
IrnI co Ien Ia trosncrtptton Iay soo
no, kIac voI Iu nhon no am cI
qua IrnI nIan doI pIan Iu ADN.
Tuy nIIen, mARN cI duoc sao
cIep Iu cac gen cIua nIung IIong
IIn can IIIeI cIo vIec Iao ra cac daI
pIan Iu proIeIn Irong Ie bao de
mang nIung IIong IIn do o dang
cac bo ba maI ma dI Iruyen den
rIbosome de dIeu kIIen qua IrnI
sInI Iong Iop proIeIn, Iay con goI
Ia qua IrnI phten no Iay djch no.
Sau kII Ioan IIanI cIuc nang cua
mnI, cIung nIanI cIong b
pIan Iuy. V vay, IIoI gIan Ion IaI

cua moI pIan Iu mARN Irong Ie
bao IIuong raI ngan.
e IIuan IIen cIo vIec IIuc
IIen cIuc nang cua mnI, mARN
Ion IaI cIu yeu o dang soI don
kIong cuon xoan pIuc Iap voI su
InI IIanI cac kIu vuc xoan kep
nIu cac IoaI ARN kIac.
Hnh IV.11. Trong cac phan t mARN
polycistron cac khu vc khong ma hoa
nam xen ke vi cac khu vc khong lam
nhiem vu ma hoa.
Trong te bao prokaryote mot si mARN duy nhat co the ma hoa cho mot hoac
mot so chuoi polypeptide. Loai mARN mang thong tin cho mot chuoi polypeptide
c goi la mARN monocistron, neu no ma hoa cho mot so chuoi polypeptide th c
goi la mARN polycistron. eukaryote phan ln mARN la monocistron. (T cistron
ay ong ngha vi t gen). Chieu dai cua mot phan t mARN tng ng vi kch
thc cua chuoi polypeptide ma no ma hoa. V du, mot chuoi polypeptide cha 100
goc aminoacid oi hoi mot mARN cha t nhat 300 goc nucleotide, v moi aminoacid
c ma hoa bi mot bo ba nucleotide. Tuy nhien, cac ban sao mARN t ADN luon
luon dai hn kch thc can thiet e ieu khien qua trnh phien ma. Phan trat t khong
trc tiep lam nhiem vu ma hoa cho trat t aminoacid cua chuoi polypeptide chnh la
trat t lam nhiem vu ieu hoa qua trnh sinh tong hp protein. Trong cac phan t
mARN polycistron cac khu vc khong ma hoa nam xen ke vi cac khu vc khong lam
nhiem vu ma hoa (hnh IV.11).
2. ARN van chuyen (tARN).
tARN co phan t tng oi nho nen viec nghien cu cau truc cua cua chung t
gap kho khan hn. Loai ARN nay chiem khoang 16% ARN tong so trong te bao, lam
nhiem vu van chuyen aminoacid trong qua trnh sinh tong hp protein. tARN cha t
75 en 90 goc nucleotide. Moi loai tARN ch tng ng vi mot aminoacid nhat nh.
Tuy nhien, mot aminoacid co the c van chuyen bi mot so tARN khac nhau. ieu
o khong co ngha la so loai tARN phai bang so codon cua moi aminoacid. V du,
glycine co 4 codon nhng ch co 3 tARN.
Mot ac iem quan trong cua tARN la
ben canh cac base chu yeu A, U, G va C trong
phan t cua chung con cha mot lng ang ke
cac base th yeu (khoang 10%). Phan ln
nhng base th yeu nay la cac dan xuat methyl
hoa va hydrogen hoa cua cac base chu yeu, ac
biet rat thng gap la acid dihyrouridilic, ky
hieu la UH
2
. Them vao o, base uracil con tao
ra mot lng nho nucleotide khong bnh thng
co ten la acid pseudouridilic, tc 5-nucleotide

cua pseudouridine, ky hieu la . Trong tARN
cung cha ca acid (ribo)thymidilic vi t cach
la mot nucleotide th yeu.
Cac base va nucleotide th yeu co tac dung lam cho phan t tARN ben vng vi
nuclease va duy tr cau truc bac ba ac trng cua phan t bang cach ngan can s hnh
thanh cau truc xoan kep nhng khu vc nhat nh ma tai o cac phan t tARN khac
nhau can gan vi mARN, ribosome va vi cac enzyme aminoacyl-tARN synthetase
ac hieu vi chung.
Nam 1955 lan au tien Holley a xac nh c hoan toan trat t nucleotide cua
tARN van chuyen alanin (tARN
ala
) gom 77 goc nucleotide. Sau o trat t nucleotide,
tc cau truc bac mot cua nhieu tARN khac cung a lan lt c xac nh. ong thi
cac nha khoa hoc cung a phat hien c rang cau truc bac mot, bac hai va bac ba cua
tat ca cac loai tARN eu co nhieu iem giong nhau (hnh IV.12,13):
- au tan cung 5 bao gi cung la pG, con au tan cung 3 la trat t pCpCpA.
Aminoacid c gan bang lien ket ester vi nhom -3OH t do cua goc acid adenylic
tan cung.
- Cac khu vc xoan kep cuc bo c bat au t base th 5 ke t au -3OH va
xen ke vi cac khu vc si n tao thanh cac thuy.
- Thuy au tien ke t au 3 c goi la thuy toan nang gom 7 nucleotide, trong
o trat t 5-pGpTppcpG-3 chung cho moi tARN. Theo nhieu tac gia, thuy nay dung
e gan vi ribosome.
- Tiep theo thuy toan nang la thuy phu vi kch thc bien ong cac loai tARN
khac nhau. Chc nang cua thuy nay cha ro.
- Sau thuy phu la thuy oi ma cha bo ba oi ma (anticodon) ac hieu cho moi
loai tARN. Boba oi ma nay luon luon nam gia mot base th yeu (thng la mot
base purine alkyl hoa) ve pha au 3 va U ve pha au 5. Trong qua trnh sinh tong
hp protein trong ribosome nh tnh bo sung gia cac bo ba oi ma cua tARN va cac
bo ba mat ma cua mARN ma hai loai ARN phoi hp c vi nhau e xac nh trat t
aminoacid cua chuoi polypeptide can c tong hp.



Hnh VI.12. S o cau truc bac hai dang la che ba cua tARN
- Tiep tuc ve pha au 5 la thuy UH
2
. Ten goi nay xuat phat t cho UH
2
luon co
mat tai ay. Trat t 3pUH
2
pGpA-5 la chung cho moi tARN. Theo quan iem c
nhieu nha khoa hoc cong nhan, thuy nay la ni tARN gan vi enzyme aminoacyl-
tARN-synthetase ac hieu.
- Cau truc bac hai vi nhng ac iem noi tren co dang mot la che ba, v vay no
thng c goi la cau truc la che ba.
- Cau truc bac ba cua tARN, theo quan iem c nhieu ngi tha nhan, co
dang ch , trong o mot au cua ch nay cha hai au tan cung 3 va 5, au oi dien
ch thuy oi ma, con goc cua no cha thuy toan nang va thuy UH
2
. Cau truc nay hnh
thanh chu yeu la nh nhng lien ket hydro khac vi cac lien ket hydro tao ra cac khu
vc xoan kep hnh IV.13).



Hnh VI.13. Mo hnh cau truc bac ba cua tARN
3. ARN ribosome (rARN).
rARN la thanh phan cau tao cua ribosome. Chung chiem hn 80% tong so ARN
cua te bao. Trong cac te bao procaryote, ma ai dien la E. coli, co ba loai rARN vi
hang so lang 23S, 16S va 5S. Hai loai 23S va 5S gop phan cung vi 34 loai protein
khac nhau, ky hieu t L1 en L34, tao nen phan di n v ribosome 50S. Trong khi
o ARN 16S cung vi 21 loai protein, ky hieu t S1 en S21, tao nen phan di n v
ribosome 30S. Khi tong hp protein, hai phan di n v 50S va 30S ket hp vi
nhau e tao nen ribosome hoat ong 70S (hnh IV.14).

Ribosome
cua te bao
eucaryote co
hang so lang
80S va c
cau tao bi hai
phan di n v
40S va 60S. Phan
di n v 40S
cha ARN 18S
ket hp vi 33
loai protein, con
phan di n v
60S cha cac loai
ARN 28S, 5,8S
va 5S ket hp vi
49 loai protein.
Cung nh
cac loai ARN
khac, chuoi
polinucleotide
cua rARN tao
nen cac khu vc
xoan kep tren c
s tnh bo sung
cua cac base nit
A-U va G-C
(hnhVI.15). Hai
phan di n v
Hnh IV.14. Cau truc khong gian cua ribosom
te bao procaryote va te bao eucaryote.

cua ribosome
phoi hp vi nhau
sao cho gia
chung co mot khe
h e si mARN
xuyen qua e
truyen at thong
tin trong qua trnh
sinh tong hp
protein. Ngoai ra,
trong ribosome
con co nhng khu
vc ac biet cho
phep chung lien
ket tam thi vi
aminoacyl-tARN
va vi si
polypeptide ang
c tong hp.
Trong s lien ket
nay rARN ong
vai tro rat quan
trong.
Hnh IV.15. rARN 16S va rARN 5S
V. PHAN GIAI ACID NUCLEIC.
1. Tac dung cuua exo- va endonuclease.
ADN va ARN b thuy phan vi s xuc tac cua cac loai nuclease va phospho-
diesterase ac hieu. Nhng enzyme nay co the c chia lam hai nhom a va b.
Nhng enzyme nhom a ch thuy phan lien ket ester gia goc phosphate vi C3 (lien
ket a trong hnh IV.7). trong khi o cac enzyme nhom b ch thuy phan lien ket ester
gia goc phosphate vi C5 (lien ket b).
Thuoc enzyme nhom a co phosphodiesterase cua noc ran. Enzyme nay thuy
phan moi lien ket a trong ADN cung nh trong ARN va giai phong nucleoside-5-
phosphate. Hoat ong cua enzyme nay oi hoi s ton tai cua nhom 3-OH t do au
tan cung mach polinucleotide. Phan ng c thc hien tng bc au tan cung nay
(hnh XII.1 a).
Thuoc nhom b co phosphodiesterase cua lach bo. No tac dung t au tan cung
cha nhom 5OH t do, thuy phan lan lt cac lien ket cua ca ADN va ARN e giai
phong nucleoside-3-phosphate (hnh XII.1 b).

Ca hai loai enzyme nay eu thuoc loai exonuclease. Khac vi chung, nhng
enzyme thuoc loai endonuclease khong oi hoi s ton tai cua nhom 3- hoac 5-OH
tan cung. Chung cong pha cac lien ket a hoac b cac v tr trong gia mach
polynucleotide. Hoat ong cua chung mang tnh chat ac hieu nhat nh oi vi thanh
phan nucleotide cua phan t ADN va ARN.
Hnh XII.1. C che tac dung cua phosphodiesterase noc ran (a) va
phosphodiesterase lach bo (b).

Deoxyribonuclease I t tuyen tuy cua bo thuy phan lien ket a gia cac
pyrimidine va purine khong phu thuoc vao v tr cua chung trong mach (vi ieu kien
mach tng oi dai). San pham la nhng oligonucleotide cha khoang 4 n v vi
goc phosphate au 5 va nhom OH au 3.
Deoxyribonuclease II t lach, tuyen c hoac vi khuan thuy phan lien ket b trong
mot so trng hp. San pham la nhng oligonucleotide cha khoang 6 n v vi goc
phosphate au 3 va nhom OH au 5.
Ribonuclease tuyen tuy ch thuy phan nhng lien ket b cua ARN ma goc
phosphate gan vi mot pyrimidine-nucleotide thong qua lien ket a (hnh XII.2).
San pham la nhng
pyrimidine-nucleoside
3-phosphate va nhng
oligo-nucleotide cha
khoang 2-5 n v
purine-nucleotide va
mot goc pyrimidine-
nucleoside ,3-
phosphate tan cung. San
pham trung gian la
nhng pyrimidine-
Hnh XII.2. C che tac dung cua ribonuclease tuyen tuy
nucleoside 2,3-phosphate vong dang goc t do hoac dang goc tan cung cua
oligonucleotide.
Ngoai nhng enzyme ke tren, ngay nay ngi ta a phat hien c nhieu loai
exonuclease va endonuclease khac t cac nguon khac nhau. Cac loai nuclease c
s dung rong rai trong viec xac nh cau truc bac mot cua acid nucleic.


2. Tac dung cua acid va kiem.
Neu thuy phan ARN bang kiem loang se hnh thanh hon hp cua 2- va 3-
nucleosidephosphate vi mot lng nho nucleoside-2,3-phosphate vong. nucleotide
vong nay la san pham trung gian va se tiep tuc b phan giai thanh nucleoside-2-
phosphate n hoac nucleoside-3-phosphate. ADN do khong cha nhom 2-OH nen
khong the tao nen nucleotide vong va v vay khong b thuy phan bang kiem.
Co the thuy phan ADN va ARN thanh base nit bang acid formic 98% 175
o
C
trong 3 phut hoac acid perchloric 12N 1OO
o
C trong mot gi. Tuy nhien, cach th
nhat lam giam lng uracil con cach th hai lam huy mot phan thymine. ADN cung co
the b thuy phan bang HCl 6N 120
o
C trong 2 gi, song mot phan purine se b mat.
Neu thuy phan ARN bang HCl 1N 100
o
C trong 1 gi, se thu c hon hp base
purine va pyrimidine-nucleotide. Nhng pyrimidine-nucleotide nay ch b thuy phan
thanh base t do trong ieu kien un soi vi acid ap suat cao trong noi ap suat hoac
ong nghiem han kn.
3. Phan giai nucleotide va nucleoside.
Nucleotide t do hnh thanh khi thuy phan acid nucleic se tiep tuc b thuy phan
thanh nucleoside nh cac enzyme 5-nucleotidase va 3-nucleotidase. Nucleosidase
se thuy phan nucleoside thanh pentose va base purine hoac base pyrimidine t do.
4. Phan giai pentose va base nit. Cac pentose tiep tuc chuyen hoa theo
con ng chuyen hoa chung cua glucide.
Base purine trong cac nhom ong vat khac nhau b phan giai vi mc o khac
nhau. ngi va mot so ong vat co vu, chim va mot so bo sat san pham cuoi cung la
acid uric. nhng loai co vu va bo sat khac cung nh giap sat acid uric tiep tuc b
phan giai thanh alantoin. Nhieu ong vat co xng song co the phan giai alantoin
thanh urea; Con nhieu loai khong xng song va thc vat lai tiep tuc phan giai urea
thanh ammoniac.
Cac giai oan chnh cua qua trnh phan giai base purine c mo ta trong hnh
XII.3.


Hnh XII.3. Phan giai base purine thanh san pham cuoi cung

Qua trnh phan giai cac base pyrimidine cha c nghien cu ay u. Cac so
lieu thu c vi sinh vat cho thay chung b phan giai thanh acid malonic, ammoniac
va CO
2
vi san pham trung gian la acid barbituric (hnh XII.4). Tng t uracil san
pham trung gian cua qua trnh d ho a thymine la acid 5-methyl-barbituric.

Hnh XII.4.Phan giai cac base pyrimidine cytosine va uracil thanh san pham cuoi cung.
Trong nhieu c the, ac biet vi khuan, cac base t do co the khong b phan giai
hoan toan ma c s dung lai e tong hp acid nucleic va cac hp chat khac. Cac
base purine t do co the biien thanh nucleotide tng ng nh
phosphoribosyltransferase:
Adenine + 5-Phosphoribosyl-1-pyrophosphate AMP + PP
vc

Guanine + 5-Phosphoribosyl-1-pyrophosphate GMP + PP
vc

Chung cung co the bien thanh nucleotide vi s xuc tac lien tiep cua hai enzyme
nucleoside phosphosylase va nucleoside kinase. V du:
Guanine + Riboso-1-phosphate Guanosine + P
vc

Guanosine + ATP GMP + ADP
Cac san pham trung gian cua qua trnh d hoa pyrimidine co the c dung e tong
hp mot so -aminoacid, v du alanine va acid -aminobutyric.


Hoa Sinh

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