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Results

This section details the preparation of pET-14b-EGFP vectors, cloning of Toehold Switch constructs, validation by DNA sequencing, and reporter expression assays for binding toehold switches to miRNAs.

Preparation of pET-14b-EGFP Vectors

Linearization of pET-14b-EGFP Plasmids by Ncol and Ndel Restriction Digestion

We amplified the purchased pET-14b-EGFP plasmids by transforming competent DH5α E. coli cells using the heat shock method. After introducing the plasmid, we grew the cells on LB plates with ampicillin to select for successful transformants. Once colonies grew, we extracted the plasmid DNA using a mini-prep procedure.

The digestion of the pET-14b-EGFP vector was performed using the following materials:

  • Plasmid DNA: 1 μg
  • 10X Restriction Buffer: 5 μL
  • Ncol-HF: 1 μL
  • Ndel: 1 μL
  • DNase-free water: add to a final volume of 50 μL
Gel Electrophoresis of Plasmids
Figure 1: Photograph of the result of gel electrophoresis showing the comparison between uncut plasmid and digested plasmid

The above gel electrophoresis demonstrates that under different forms of the plasmid, the plasmids travelled at different speeds under gel electrophoresis. This is essential for separating the digested plasmid from the undigested ones. It is because when digested, the plasmid would change from its supercoiled form, which travelled faster in gel due to higher compactness, to its linearised form which travelled slower in gel.

Cloning of Toehold Switch Constructs

Transformation of Recombinant Plasmids

Unsuccessful Attempts

Attempt 1: After ligating the pET-14b-EGFP plasmid with Toehold Switch 1 and Toehold Switch 2, we transformed E. coli with the recombinant plasmids: Toehold Switch 1 plasmids (T1 plasmids) and Toehold Switch 2 plasmids (T2 plasmids). The bacteria were then cultured on agar plates containing LB medium and ampicillin. We also set up two controls: a positive control with bacteria transformed with the pET-14b-EGFP plasmid (non-recombinant) and a negative control with no bacteria.

Results

Agar plate type
Colonies of bacteria Observed?
1. No Bacteria (Negative control)
No
2. Bacteria transformed with non-recombinant Plasmids (Positive Control)
Yes (many colonies)
3. Bacteria transformed with T1 Plasmids
1 colony
4. Bacteria transformed with T2 Plasmids
1 colony
Transformation Attempt 1
Figure 2: Photograph of the result of transformation attempt 1

Interpretation and conclusion

The result of negative control indicated that the plate and antibiotic were functioning correctly, without any contamination. The positive control confirmed that the transformation procedure was successful, as the bacteria expressed the antibiotic resistance gene, allowing them to survive on the ampicillin plates.

We observed one colony of bacteria on each agar plate transformed with T1 and T2 plasmids, suggesting that the transformation of bacteria with the Toehold Switch plasmids may have been successful. Since ampicillin selection alone cannot confirm the presence of recombinant clones, we picked the colonies and sent them for DNA sequencing.

However, the DNA sequencing results showed that both Toehold Switches 1 and 2 were not ligated into the plasmids; the base sequences of the plasmids remained unchanged, and we could not find the sequences of Toehold Switches 1 and 2 between NcoI and NdeI.

Due to limitations in our laboratory skills for gel extraction and plasmid purification, we decided to make another attempt.

Attempt 2: We restarted the experiment with some modifications: we extracted as much gel as possible, aimed to maximize the concentration of the plasmids and proper mixing of inserts into vectors. We then carried out the transformation of bacteria with the T1 and T2 plasmids once again. The results are shown in the following figures.

Results

Agar plate type
Colonies of bacteria Observed?
1. No Bacteria (Negative control)
No
2. Bacteria transformed with non-recombinant Plasmids (Positive Control)
Yes (many colonies)
3. Bacteria transformed with T1 Plasmids
No
4. Bacteria transformed with T2 Plasmids
1 colony
Transformation Attempt 2 T1
Figure 3: Photograph of the transformation result of Toehold Plasmid 1 in the second attempt
Transformation Attempt 2 T2
Figure 4: Photograph of the transformation result of Toehold Plasmid 2 in the second attempt

Interpretation and conclusion

There was one colony of bacteria transformed with T2 plasmids on the agar plate. We picked the colony and sent it to DNA sequencing. Unfortunately, the sequencing result showed that the insertion of Toehold Switch 2 into the plasmid failed, as the base sequence of the plasmid remained unchanged and the base sequences of Toehold switch 2 cannot be found between NcoI and NdeI. The result of DNA sequencing shows that the ligation was unsuccessful. This can be caused by the low concentration of plasmids (vectors) or the ratio of insert to vector.

Attempt 3: This time, we used maxiprep instead of miniprep to obtain a higher concentration of plasmid (the vector). We added the 5’ phosphate group to oligos of the Toehold Switches for insertion. Then we carried out ligation and transformation again. To increase our chances of success, we used another ratio of concentration of the plasmid (the vector) to the Toehold Switches 1 and 2 (the inserts), which was 1:10. After ligation, we did the transformation. Then we cultured the transformed bacteria in LB medium with ampicillin.

Results

Agar plate type
Colonies of bacteria Observed?
1. No Bacteria (Negative control)
No
2. Bacteria transformed with non-recombinant Plasmids (Positive Control)
Yes (many colonies)
3. Bacteria transformed with T1 Plasmids (vector: insert in 1:10)
2 colonies
5. Bacteria transformed with T2 Plasmids (vector: insert in 1:10)
No
Transformation Attempt 3 Transformation Attempt 3 Transformation Attempt 3 Transformation Attempt 3
Figure 5: Photographs of the transformation result of toehold plasmid 1 and 2 in the third attempt

Interpretation and conclusion

Only bacteria transformed with T1 Plasmids could be cultured. Bacteria were picked up and were sent for DNA sequencing.

However, the result of DNA sequencing showed that the Toehold Switch 1 failed to be inserted into the plasmids. We had to revise our protocol.

Successful Cloning

We used a high insert-to-vector ratio (20:1) to ligate toehold switches 7, 8, and 9 into pET-14b-EGFP plasmids extracted via maxiprep. The resulting recombinant plasmids were transformed into E. coli and plated on LB agar containing ampicillin for colony growth.

Results:

Agar plate
E.coli colonies
1. Negative control: E.coli transformed with cut (linearised) plasmid vector
0
2. Positive control: E.coli transformed with plasmid vector without insert
>100
3. E.coli transformed with T7 recombinant plasmids
0
4. E.coli transformed with T8 recombinant plasmids
>100
5. E.coli transformed with T9 recombinant plasmids
0
Transformation results of toehold switches 7, 8 and 9 plasmids on agar plates
Fig. 6. Transformation results of toehold switches 7, 8 and 9 plasmids on agar plates.

Interpretation and conclusion

The negative control verified that the antibiotic was effective and that no contamination occurred during bacterial culture.

The positive control confirmed that E. coli transformed with plasmid vectors with intact ampicillin resistance gene could successfully grow.

Colonies observed on agar plates with E. coli transformed with T8 plasmids suggested cloning of T8 may be successful. However, since ampicillin selection alone cannot confirm recombinant clones, we verified the T8 recombinant plasmids through DNA sequencing.

Validation by DNA Sequencing

The DNA sequencing results confirmed that our E.coli colonies were actually transformed with our toehold switch 8 recombinant plasmids, proving our success in cloning.

Sequencing report from
Figure 7: Sequencing report of toehold 1
Sequencing report of toehold 1
Figure 8: Sequencing report of toehold 2
Sequencing report of toehold 2
Figure 9: Sequencing report of toehold 3
Sequencing report of toehold 3
Figure 10: Sequencing report of toehold 7
Sequencing report of toehold 7
Figure 11: Sequencing report of toehold 8
Sequencing report of toehold 8
Figure 12: Sequencing report of toehold 9

Reporter Expression

Binding of toehold switches 1,2,3 to miRNA-92a-3p; toehold switches 7,8,9 to miRNA-135b-5p

Attempt 1 (Preliminary test): Cell free expression of Toehold Switch 3 plasmids only

To test the binding of toehold switches, we used an in vitro expression (IVE) kit, which allows the transcription and translation of the EGFP gene on plasmids in a cell-free system. We mixed 1.5 µL of 5 µM miRNA samples with 62.5 ng of plasmid, with or without the toehold switches, in a total reaction volume of 6 µL. The samples were then observed under blue light. Green fluorescence would indicate successful binding of the miRNA to the toehold switches, leading to the opening of the hairpin structure and subsequent expression of EGFP.

A negative control, containing no plasmid, and a positive control, containing the non-recombinant plasmid, were also included. The table below shows the contents of the tubes in the IVE kit and shows the contents of 6 tubes in in vitro expression.

Tube
Presence of miRNA
Contents
Role
1
No
No plasmid
Negative control
2
No
T3 plasmid
Experimental
3
No
Plasmid
Positive control
4
Yes
No plasmid
Negative control
5
Yes
T3 plasmid
Experimental
6
Yes
Plasmid
Positive control
The tubes under blue light
Figure 13: The tubes under blue light

As minimal differences were found between the positive and negative controls under influence of blue light, we found that it was necessary to use a plate reader for measuring the fluorescence intensity. Also, we had to optimize the amount of plasmids we used in the in vitro expression kit.

Optimisation of Positive Control

In order to optimize the amount of plasmids used in the in vitro expression kit, a total of 3 trials have been performed.

Trial 1

Content of the tubes in IVE Kit

Tube No.
A
B
C
D
E
Solution A (µL)
2.0
2.0
2.0
2.0
2.0
Solution B (µL)
1.5
1.5
1.5
1.5
1.5
pET-14b-EGFP (plasmid) (µL)
500 ng 1.5
300 ng 1.5
187.5 ng 1.5
37.5 ng 1.5
7.5 ng 1.5
Total (µL)
5.0
5.0
5.0
5.0
5.0
The tubes under blue light
Figure 14: The tubes under blue light

Measurement of fluorescence intensity in tubes with different amount of plasmids

Samples
Blank
100 µL PBS
A (500 ng plasmid)
B (300 ng plasmid)
C (187.5 ng plasmid)
D (37.5 ng plsamid )
E (7.5 ng plasmid)
Fluorescence intensity
0
56
1194
1308
1209
1205
2128
Optimisation: Trial 2 and 3
The tubes of different amounts of plasmids under blue light
Figure 15: The tubes of different amount of plasmids under blue light: The tube order (from left to right): First row: A (500 ng), B(300ng), C(187.5ng), D(37.5ng), E(7.5ng), E*(7.5ng) for Trial 3. Second row: A (500 ng), B(300ng), C(187.5ng), D(37.5ng), E(7.5ng),E* (human error involved in this set up) for Trial 2.

Trial 2

Measurement of fluorescence intensity in tubes with different amount of plasmids

Samples
Blank
100 µL PBS
A (500 ng plasmid)
B (300 ng plasmid)
C (187.5 ng plasmid)
D (37.5 ng plsamid)
E (7.5 ng plasmid)
Fluorescence intensity
0
-3.0
-77.0
946.0
922.0
977.0
1341.0
The tubes of different amounts of plasmids under blue light
Figure 16: The tubes of different amount of plasmids under blue light: From left to right: A, B, C, D, E

Trial 3

Measurement of fluorescence intensity in tubes with different amount of plasmids

Samples
Blank
100 µL PBS
A (500 ng plasmid)
B (300 ng plasmid)
C (187.5 ng plasmid)
D (37.5 ng plasmid)
E (7.5 ng plasmid)
E* (7.5 ng plasmid)
Fluorescence intensity
0
-3.0
1287.0
1187.0
2019.0
965.0
844.0
723.0
The tubes of different amount of plasmids under blue light Trial 2
Figure 17:The tubes of different amount of plasmids under blue light: From left to right: A, B, C, D, E, E* (human error involved)

Average fluorescence intensity in tubes with different amount of plasmids

Blank 100µL PBS
500 ng Plasmid
300 ng Plasmid
187.5ng Plasmid
37.5 ng Plasmid
7.5 ng Plasmid
Trial 1
0 56
1194
1308
1209
1205
2128
Trial 2
0 -3
-77
946
922
977
1341
Trial 3
0 -3
1287
1187
2019
965
844
Average
0 16.7
801.3
1147.0
1383.3
1049.0
1437.7
The graph shows the fluorescence expression by different amounts of plasmids in the IVE kit
Figure 18. The graph shows the fluorescence expression by different amounts of plasmids in the IVE kit

When comparing the fluorescence intensity across different amounts of plasmids, the highest fluorescence was observed with 7.5 ng of plasmids. Therefore, 7.5 ng was identified as the optimal amount for subsequent experiments on in vitro expression.

Attempt 2: Cell free expression of Toehold Switch plasmids

The table below shows the contents of each tube in the IVE kit. Tubes 1 to 5 did not contain any target miRNA, while tubes 6 to 10 included 5 µM of the target miRNA (miRNA-92a-3p). The negative control was the tube without any plasmids, whereas the positive control contained the plasmid pET-14b-EGFP.

Content of the tubes used in in vitro Expression

Tube No.
1
2
3
4
5
Nuclease-free water
Toehold Switch 1 plasmid
Toehold Switch 2 plasmid
Toehold Switch 3 plasmid
Plasmid (pET-14b-EGFP)
-miRNA
-miRNA
-miRNA
-miRNA
-miRNA
Tube No.
6
7
8
9
10
Nuclease-free water
Toehold Switch 1 plasmid
Toehold Switch 2 plasmid
Toehold Switch 3 plasmid
Plasmid (pET-14b-EGFP)
+miRNA
+miRNA
+miRNA
+miRNA
+miRNA

Tubes 1 and 6 serve as the negative controls: These ensure that any observed fluorescence results specifically from the interaction of the miRNA with the toehold switches, rather than from other sources. Since no plasmid is present, any fluorescence in these controls suggests possible contamination or nonspecific reactions occurring within the kit.

Tubes 5 and 10 serve as the positive controls: They confirm that the experimental conditions are suitable for the desired reaction. With the non-recombinant plasmid present, the observation of green fluorescence indicates that the components of the IVE kit and the overall experimental setup are functioning correctly.

The table below shows the fluorescence intensity of each tube in in vitro expression.

Sample
Fluorescence intensity (without miRNA added)
Fluorescence intensity (with miRNA added)
PBS*
131.5
131.5
Nuclease free water (negative control)
1899.5
1244.5
Toehold Switch 1 plasmid
2195.5
4517.5
Toehold Switch 2 plasmid
4992.5
11074.5
Toehold Switch 3 plasmid
4497.5
2504.5
pET-14b-EGFP plasmid (positive control)
47089.5
12585.5

*PBS refers to the tube with Phosphate-buffered saline (PBS) only. PBS is a buffer solution (pH ~ 7.4) commonly used in biological research. All the samples were made up to 100 µL by adding PBS before measurement of fluorescence, so It serves for measuring the background fluorescence.

Fluorescence intensity expressed by different samples
Figure 19: Fluorescence intensity expressed by different samples

Based on the data, the fold change of fluorescence intensity (+miRNA / -miRNA) was calculated and presented in the table below.

Sample
Fold Change of fluorescence intensity (+miRNA / -miRNA)
Toehold Switch 1 plasmid
2.12
Toehold Switch 2 plasmid
2.25
Toehold Switch 3 plasmid
0.54
Fold change in fluorescence intensity expressed by Toehold Switch plasmids with miRNA added compared to without miRNA added
Figure 20: Fold change in fluorescence intensity expressed by Toehold Switch plasmids with miRNA added compared to without miRNA added

From the above data, we can see that: Toehold Switch 1 displayed a fold change of 2.12 in fluorescence, indicating an increase upon miRNA addition. This suggests that it can effectively detect miRNA, though it shows slightly less efficacy compared to Toehold Switch 2. ⁠Toehold Switch 2 achieved a fold change of 2.25, representing the highest increase in fluorescence among the tested switches for miR-92a-3p. This reinforces its designation as the most effective candidate for miR-92a-3p detection.

Toehold Switch 3 showed a decrease in fluorescence, likely due to issues with binding or structural stability that prevent effective translation. This indicates that it is unsuitable for miRNA detection.

Negative controls (nuclease-free water and setups without Toehold Switches) confirmed that the fluorescence change was due to the interaction of miRNA with the toehold switches, rather than non-specific effects.

Conclusion

Both Toehold Switch 1 and Toehold Switch 2 exhibited sufficient fluorescence after the addition of miRNA, but Toehold Switch 2 was slightly more effective, showing a higher fold change than that of Toehold Switch 1.

While both switches are viable candidates, the slight advantage in Toehold Switch 2 makes it the preferred choice for further investigation.