2 주차 molecular and biological chemistry 2. reading frame frame 모든 진핵생물은 + strand...
TRANSCRIPT
![Page 1: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/1.jpg)
2 주차Molecular and
Biological Chemistry 2
![Page 2: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/2.jpg)
Reading frame
Frame
• 모든 진핵생물은 + strand 에만 정보 갖고 있음• 원핵생물은 – strand 에도 정보를 갖고 있지만 +strand 와 서로 겹치지
않음• Virus 는 +, - 모두를 사용하며 , 사용구역이 겹치기도 하고 , 다른 frame
을 사용하기도 함 .
![Page 3: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/3.jpg)
Amborella (eukaryotes) uses only + strand without over-lap!
http://amborella.org
![Page 4: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/4.jpg)
Magnolia kobus DC.chloroplast
genome159,245bp
Magnolia chloroplast uses + and – strand without overlap
![Page 5: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/5.jpg)
Cyanobacterium also uses both strand without overlap
Cyanobacteriumcompletegenome
![Page 6: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/6.jpg)
Herpes simplex virus
DNA virus uses both strand and multiple frame!!!
![Page 7: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/7.jpg)
Ethidiumbromide (EtBr): reagent used for DNA detection. It causes frame shift mu-tation (carcinogen)!
Frame shift mutation
![Page 8: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/8.jpg)
Polymerase Chain Reaction (PCR)
• A technique for the in vitro amplification of specific DNA se-quences by the simultaneous primer extension of comple-mentary strands of DNA (>105).
• PCR method was devised and named by Mullis and col-leagues at the Cetus Corporation (Mullis and Faloona, 1987)
• The principle had been described in detail by Khorana et al. (Kleppe et al., 1971) over a decade earlier.
• Kary Mullis received the Novel prize (1993).
PCR
![Page 9: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/9.jpg)
3) DNA POLYMERASE
GT T T TC CA A G G G
2) PRIMER3’5’
AC TA C G TA CG TA C GT G T C CT AC G G TT CT T CT T A G T TG T A C CA CA G TA C TCG TA C T CG TA C T AAA
1) TEMPLETE3’ 5’
…
4) dNTP’s pool
G
C
A
T
dATP
dCTP
dGTPdTTP
5) Mg++
G
CA
T
A
A
A
A
C
C
C
C
G
T
G
T
G
T G
T
GT T T TC CA A G G G
AC TA C G TA CG TA C GT G T C CT AC G G TT CT T CT T A G T TG T A C CA CA G TA C TCG TA C T CG TA C T AAA
3’
5’
5’
…
Newly synthesised strand
C A A G G A CG T C CA A A T G C A A G A A A A A A AG G G G GT T T T T TC C C CA AG GAA
G
T
DNA polymerization: DNA polymerase extends a primer by using a complementary strand as a template.
![Page 10: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/10.jpg)
Schematic diagram of PCR
… …Denature
94 ℃3’
5’
5’… …
3’
3’
5’
5’
…
…
…
…
Anneal primers
ca. 50 ℃5’3’
3’
5’…
3’
GenomicDNA
…5’
3’
… …
…5’ 3’
…3’ 5’
…
5’ 3’
…3’ 5’
…Synthesize
Newstrand72 ℃
…
3’
5’
5’
…
…
…
…
5’3’
3’
5’
…5’ 3’
Anneal primers
ca. 50 ℃
5’
…3’
5’
5’3’5’ 3’
3’
5’
5’
SynthesizeNew
strand72 ℃
3’ 5’
…
3’
…5’ 3’
… …
5’ 3’
3’ 5’
……
5’
…
3’…
3’ 5’
3’ 5’… …
5’… …3’
3’ 5’
…5’ 3’Denature
94 ℃
…
![Page 11: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/11.jpg)
…
… …
……
…
…
…
Denature94 ℃
…
… …
…
…
…
… …
Anneal primers
ca. 50 ℃
…
… …
…
…
…
… …
…
…
SynthesizeNew
strand72 ℃
…
![Page 12: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/12.jpg)
![Page 13: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/13.jpg)
![Page 14: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/14.jpg)
심해 열수구(hydrothermal vents)
![Page 15: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/15.jpg)
Taq. DNA Polymerase:
• Most commonly used DNA polymerase for PCR Isolated from Thermus aquaticus Heat stable polymerase
• PCR technique put to practical use by finding of Taq.
• The use of a heat stable enzyme has two major advantages:
1) replenishment after each heating step is not required, thus simplifying the process
2) the enzyme is active at high temperatures, where anneal-ing of the oligonucleotide primers is more specific and DNA synthesis more rapid.
![Page 16: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/16.jpg)
Thermophilic DNA poly-merases
Thermus aquaticus (Taq) Thermus thermophilus (Tth) Bacillus stearothermophilus (Bst) Pyrococcus furiosis (Pfu)
![Page 17: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/17.jpg)
Reaction components
1) Thermophilic DNA polymerases
• Thermus aquaticus (Taq) Thermus thermophilus (Tth) Bacillus stearothermophilus (Bst) Pyrococcus furiosis (Pfu)
• Genetically modified for improving proof reading function (3’ to 5’ exonuclease function) and for hot start PCR.
![Page 18: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/18.jpg)
TA cloning
![Page 19: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/19.jpg)
Reaction components
1) Thermophilic DNA polymerases
• Thermus aquaticus (Taq) Thermus thermophilus (Tth) Bacillus stearothermophilus (Bst) Pyrococcus furiosis (Pfu)
• Genetically modified for improving proof reading function (3’ to 5’ exonuclease function) and for hot start PCR.
![Page 20: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/20.jpg)
Final-extend72 ℃
An example of a PCR method
0
20
40
60
80
Temp. ℃
100
Holdprogram
Pre-denature95 ℃
Anneal55 ℃
Extend72 ℃
Cycleprogram
Holdprogram
Cycle 1 Cycle 2 • • • Cycle 30
Denature95 ℃
Soak4 ℃
![Page 21: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/21.jpg)
Reaction components
2) Deoxynucleotide Triphosphates
• Low dNTP concentrations minimize mispriming at nontarget sites
and reduce the likelihood of extending misincorporated nucleotides (Innis et al. 1988)
![Page 22: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/22.jpg)
Reaction components
3) Primers
• Generally, use over 18 nucleotides
• Genome size of higher plants: 5X108-6X109
• Check list for primer design: 1) Similar Tm values are recommended for two primers 2) Avoid self-complement sequences 3) Avoid primer dimer formation
Probability of presence of same nucleotide sequences
6 mer 1/46= 1 / 4X103
9 mer 1/49= 1 / 2.6X105
12 mer 1/412= 1 / 1.7X107
15 mer 1/415= 1 / 1.1X109
18 mer 1/418= 1 / 6.9X1010
21 mer 1/421= 1 / 4.4X1012
Q: 왜 PCR 을 할 때 사용하는 primer 는 주로 18bp 이상을 사용해야 할까 ?
![Page 23: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/23.jpg)
![Page 24: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/24.jpg)
![Page 25: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/25.jpg)
![Page 26: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/26.jpg)
![Page 28: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/28.jpg)
Primer order 의 예
![Page 29: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/29.jpg)
Reaction components
4) Reaction buffer
• Mainly modified from Saiki et al. (1988)
• Low conc. of Mg++: reaction failed High conc. of Mg++: mis-pairing
pseudo bands
Components of PCR buffer
Saiki et al. (1988): Suh lab: Tris pH 8.4 10mM 20mM KCl 50mM 50mM MgCl2 1.5mM 1.5mM gelatin 0.01% - NP40 0.01% - Tween-20 0.01% 0.001%
![Page 30: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/30.jpg)
Reaction components
5) Template DNA
• General amount of template DNA: 105-106 target molecules
• 1 ug of human DNA 10 ng of yeast DNA ≒ 3X105 molecules of single copy gene 1 ng of E.coli DNA
• Quantification of extracted DNA: 1) spectrophotometer: OD260
2) spot test in agarose gel
![Page 31: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/31.jpg)
UV-spec vs. Nanodrop
An A260 of 1.0 = 50 mg/ml concentration of DNA.
![Page 32: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/32.jpg)
Gel electrophoresis
침강제 +Dye+DNA
![Page 33: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/33.jpg)
An Example of band (spot) test
• Marker: PCR Marker (Promega G3161)
- Marker concentration: 5 ul contains 30-40 ng of
each DNA fragments
- Sample DNA concentration: 6-8 ng/ul
∴ dilute to 1/10 use 2 ul of DNA for 100ul reaction (1.2-1.6 ng)
![Page 34: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/34.jpg)
Thermal condition and cycle number
e.g. PCR amplification of ndhF gene in Magnoliaceae (Kim et al., 2001)
Pre-denaturation: 95 ℃ 10 min ( AmpliTaq. Gold)
Denaturation: 95 ℃ 1 min Primer annealing: 55 ℃ 1 min 30 cycles Primer extension: 72 ℃ 1 min
Final extension: 72℃ 7 min
PCR 온도 조건의 예
![Page 35: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/35.jpg)
Final-extend72 ℃
An example of a PCR method
0
20
40
60
80
Temp. ℃
100
Holdprogram
Pre-denature95 ℃
Anneal55 ℃
Extend72 ℃
Cycleprogram
Holdprogram
Cycle 1 Cycle 2 • • • Cycle 30
Denature95 ℃
Soak4 ℃
![Page 36: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/36.jpg)
![Page 37: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/37.jpg)
Depending on reaction conditions and thermal cycling, one or more of the following may influence plateau:
1) Utilization of substrates (dNTPs or primers) 2) Stability of reactants (dNTPs or enzymes) 3) End-product inhibition (pyrophosphate, duplex DNA)…
![Page 38: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/38.jpg)
Non-PCR amplification
• Cloning - PCR product 가 heterogenous 할 때 각각의 염기서열을 결정하려면 cloning 이 필요함 .• RCA (Rolling circle amplificaion)
![Page 39: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/39.jpg)
• Cloning
![Page 40: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/40.jpg)
![Page 41: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/41.jpg)
• RCA (Rolling circle amplificaion) used for small amount of template such as DNA in
herbarium sheets or even fossils.
![Page 42: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/42.jpg)
Krause et al. 2006. Multiplification of the mammoth mitochondial genome and the evolution of Elephantidae. Nature 439: 724-727.
Poinar et al. 2006 Metagenomics to paleogenomics: large-scale sequencing of mammoth DNA. Science 311: 392-394.
![Page 43: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/43.jpg)
1993
최신 DNA 추출기술과 PCR 기술의 발전은 DNA 가 매우 적게 남아 있는 화석에서 DNA 염기서열 결정을
가능하게 함
![Page 44: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/44.jpg)
![Page 45: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/45.jpg)
![Page 46: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/46.jpg)
![Page 47: 2 주차 Molecular and Biological Chemistry 2. Reading frame Frame 모든 진핵생물은 + strand 에만 정보 갖고 있음 원핵생물은 – strand 에도 정보를 갖고 있지만](https://reader036.vdocuments.pub/reader036/viewer/2022062409/56649cad5503460f9497001a/html5/thumbnails/47.jpg)
Magnolia latahensis (Berry) Brown