DNA isolation from plant tissue is a fundamental technique in plant molecular biology, genetics, taxonomy, and biotechnology, providing genomic DNA for various molecular and genetic analyses.
Plant DNA extraction can be more challenging than DNA isolation from animal cells because plant cells contain rigid cellulose- and hemicellulose-rich cell walls, along with high levels of polysaccharides, polyphenols, and other secondary metabolites. These compounds can co-purify with DNA, reduce its purity, inhibit enzymatic reactions, and interfere with downstream applications such as PCR, DNA sequencing, and restriction digestion.
Therefore, plant DNA extraction requires robust and selective methods that efficiently disrupt plant tissues while separating genomic DNA from proteins, polysaccharides, polyphenols, pigments, and other cellular contaminants.
The cetyltrimethylammonium bromide (CTAB) method is one of the most widely used and reliable approaches for isolating genomic DNA from plant tissues. Originally described by Doyle and Doyle (1987), the CTAB protocol has become a classic method because it is effective across a wide range of plant species, including those containing high levels of secondary metabolites.
The CTAB method is based on the selective separation of nucleic acids under high-salt conditions. CTAB facilitates the separation of DNA from cellular components, while high salt concentrations help minimize the co-precipitation of polysaccharides. Subsequent organic extraction steps help remove proteins, lipids, pigments, and other contaminants, resulting in DNA suitable for downstream molecular applications.
Numerous modifications of the CTAB protocol have been developed to improve DNA yield and purity, particularly for challenging materials such as silica-dried leaves, herbarium specimens, and plant tissues rich in phenolic compounds (Sahu et al., 2016; Schenk et al., 2023a; Schenk et al., 2023b).
Commercial and academic protocols, including those described by Zymo Research (2022) and Biotech Beacon (2025), have further adapted the CTAB approach for routine laboratory applications.
Overall, CTAB-based extraction remains a robust, adaptable, and cost-effective method for plant genomic DNA isolation, particularly when high-quality DNA is required for PCR, sequencing, restriction digestion, genetic analysis, and other molecular biology applications.
Key Reagents of DNA Isolation from Plant Tissues
The CTAB-based plant DNA isolation method uses a combination of detergents, buffering agents, chelating agents, salts, reducing agents, organic solvents, alcohols, and optional enzymes. Each reagent has a specific biochemical role in cell lysis, DNA stabilization, removal of contaminants, and DNA precipitation.
CTAB (Cetyltrimethylammonium bromide) — 2% (w/v): Acts as a cationic detergent that disrupts cellular and nuclear membranes and facilitates the separation of DNA from polysaccharides and other cellular components.
Tris-HCl — 100 mM, pH 8.0: Maintains a stable and slightly alkaline pH, helping preserve DNA integrity during the extraction process.
EDTA (Ethylenediaminetetraacetic acid) — 20 mM: Chelates divalent metal ions such as Mg²⁺ and Ca²⁺, which are required by many nucleases. This helps inhibit DNase activity and protects genomic DNA from degradation.
NaCl — 1.4–2.5 M: Provides high ionic strength and promotes the separation of DNA from polysaccharides, helping reduce polysaccharide contamination in the DNA preparation.
β-Mercaptoethanol — 0.2–1% (v/v): Functions as a reducing agent and helps limit the oxidation of phenolic compounds. This is particularly important when extracting DNA from plants with high levels of polyphenols.
Chloroform:Isoamyl alcohol — 24:1 (v/v): Used during organic extraction to facilitate the removal of proteins, lipids, pigments, and other hydrophobic contaminants. Isoamyl alcohol also helps reduce foaming and improves phase separation.
Isopropanol — approximately equal volume: Used to precipitate DNA from the aqueous phase. DNA has reduced solubility in the presence of isopropanol and salt, allowing it to be recovered as a precipitate.
70% Ethanol: Used to wash the DNA pellet and remove residual salts and other soluble contaminants while retaining the precipitated DNA.
RNase A — optional: Added when RNA contamination needs to be minimized. RNase A enzymatically degrades RNA, improving the purity of the isolated DNA.
The exact concentrations and volumes may vary depending on the plant species, tissue type, sample condition, and specific CTAB protocol being used. These reagents and their functions are adapted from established plant DNA extraction procedures (Doyle & Doyle, 1987; Zymo Research, 2022; Biotech Beacon, 2025).
Principle of DNA Isolation from Plant Tissues
The CTAB method is based on the differential solubility and separation of nucleic acids and cellular contaminants under detergent, high-salt, and organic extraction conditions. The overall process involves cell disruption, membrane lysis, removal of contaminants, DNA precipitation, and purification.
Cell disruption and lysis: Plant tissue is mechanically disrupted and treated with CTAB extraction buffer to break down cell walls and cellular membranes. CTAB, a cationic detergent, solubilizes membrane lipids and helps release DNA and other cellular components into the extraction mixture.
Separation of polysaccharides: Under high-salt conditions provided by NaCl, CTAB interacts with acidic polysaccharides and promotes their separation from DNA. This is particularly important because polysaccharide contamination can interfere with DNA purity and downstream molecular applications (Doyle & Doyle, 1987).
Protection of DNA: EDTA chelates divalent metal ions such as Mg²⁺ and Ca²⁺ that are required for the activity of many nucleases, thereby helping protect DNA from enzymatic degradation. Tris-HCl maintains a stable pH during extraction, supporting DNA stability.
Removal of phenolic compounds: β-Mercaptoethanol acts as a reducing agent and helps prevent the oxidation of phenolic compounds. Oxidized phenolics can interact with and become associated with DNA, resulting in reduced DNA purity and potentially interfering with downstream applications (Sahu et al., 2016).
Organic extraction: Following cell lysis, chloroform:isoamyl alcohol is used to separate proteins, lipids, pigments, and other cellular debris from the nucleic acid-containing aqueous phase. After centrifugation, DNA remains predominantly in the aqueous phase, while many contaminants partition into the organic phase or form an interfacial layer.
DNA precipitation: DNA is recovered from the aqueous phase by adding isopropanol or another suitable alcohol. Because DNA has lower solubility in alcohol, particularly in the presence of salt, it precipitates and can be collected by centrifugation.
DNA washing: The resulting DNA pellet is washed with 70% ethanol to remove residual salts, CTAB, and other soluble contaminants while keeping the DNA precipitated.
Final DNA recovery: After removal of the wash solution and appropriate drying, the purified DNA is resuspended in a suitable buffer or nuclease-free water. The resulting genomic DNA can then be used for downstream applications such as PCR, restriction digestion, sequencing, genotyping, and other molecular analyses.
Protocol of DNA Extraction from Plant Tissues (CTAB Method)
Sample Preparation
Weigh approximately 50–100 mg of plant tissue, using fresh, silica-dried, or appropriately preserved material.
Clean the tissue to remove surface contaminants and excess moisture.
Freeze the sample thoroughly using liquid nitrogen.
Grind the frozen tissue in a pre-chilled mortar and pestle until a fine, homogeneous powder is obtained.
Immediately transfer the powdered tissue into a sterile 1.5 mL microcentrifuge tube.
Add 700 µL of pre-warmed CTAB extraction buffer containing:
2% (w/v) CTAB
100 mM Tris-HCl, pH 8.0
20 mM EDTA
1.4–2.5 M NaCl
Add 5–10 µL β-mercaptoethanol freshly before use to achieve approximately 0.2–1% final concentration.
Mix gently by inverting the tube to ensure that the powdered tissue is completely suspended.
Cell Lysis
Incubate the tube at 60–65°C for 30–60 minutes.
Gently invert the tube approximately every 10 minutes to promote uniform lysis and mixing.
During incubation:
CTAB disrupts cellular and nuclear membranes and facilitates the release of DNA.
High NaCl concentration promotes the separation of DNA from polysaccharides.
β-Mercaptoethanol helps prevent oxidation of phenolic compounds.
After incubation, allow the lysate to cool to room temperature for approximately 5 minutes before proceeding to organic extraction.
Chloroform–Isoamyl Alcohol Extraction
Add 700 µL chloroform:isoamyl alcohol (24:1, v/v) to the lysate.
Mix gently by inversion for 5–10 minutes. Avoid vigorous vortexing, which can shear high-molecular-weight genomic DNA.
Centrifuge at 12,000–16,000 × g for 10 minutes at room temperature.
Following centrifugation, three distinct regions may be observed:
Lower organic phase: Primarily chloroform containing extracted lipids and other hydrophobic contaminants.
Interphase: Enriched with denatured proteins and cellular debris.
Upper aqueous phase: Contains the extracted DNA.
Carefully transfer approximately 500–600 µL of the upper aqueous phase into a fresh sterile microcentrifuge tube without disturbing the interphase.
If the interphase remains visibly cloudy or contaminated, repeat the chloroform:isoamyl alcohol extraction before DNA precipitation.
RNA Removal (Optional)
If RNA contamination needs to be reduced, add 5 µL RNase A (10 mg/mL) to the recovered aqueous phase.
Mix gently by inversion.
Incubate at 37°C for 30–45 minutes.
Briefly centrifuge the tube to collect any condensation from the tube walls.
DNA Precipitation
Add approximately 0.6–1.0 volume of chilled isopropanol to the aqueous phase. For 500 µL of aqueous solution, this corresponds to approximately 300–500 µL isopropanol.
Gently invert the tube 10–15 times until the solution becomes cloudy and DNA precipitation is apparent.
Incubate at −20°C for 20–30 minutes to enhance DNA precipitation.
Centrifuge at 12,000–16,000 × g for 10–15 minutes at 4°C.
Carefully remove and discard the supernatant without disturbing the DNA pellet, which may appear white, translucent, or only faintly visible depending on DNA yield.
DNA Recovery and Washing
Add 1 mL of chilled 70% ethanol to the DNA pellet.
Gently invert the tube to wash the pellet and remove residual salts and other contaminants.
Centrifuge at approximately 10,000 × g for 5 minutes at 4°C.
Carefully discard the ethanol without dislodging the pellet.
If substantial salt contamination is suspected, perform a second 70% ethanol wash.
Allow the pellet to air-dry at room temperature for approximately 5–10 minutes. Avoid excessive drying because an over-dried DNA pellet can be difficult to dissolve.
DNA Resuspension and Storage
Resuspend the DNA pellet in approximately 30–50 µL of either:
TE buffer: 10 mM Tris-HCl, 1 mM EDTA, pH 8.0, or
Nuclease-free water
Allow the DNA to dissolve at room temperature for 15–30 minutes. For difficult-to-dissolve pellets, incubation at 4°C overnight can facilitate complete resuspension.
Mix gently rather than vigorously vortexing to minimize mechanical shearing of genomic DNA.
Store the purified DNA under appropriate conditions, typically at 4°C for short-term use or −20°C for longer-term storage.
The extracted DNA can subsequently be evaluated for concentration, purity, and integrity before use in downstream applications such as PCR, sequencing, restriction digestion, and genotyping.
Observations and Results
DNA pellet formation: Following isopropanol precipitation and centrifugation, successfully extracted genomic DNA typically appears as a white, translucent, or slightly cloudy pellet at the bottom or side of the microcentrifuge tube. The visibility of the pellet depends on the amount of DNA recovered.
DNA integrity: When analyzed by agarose gel electrophoresis, high-quality genomic DNA generally appears as a strong, high-molecular-weight band with minimal smearing, indicating limited DNA degradation. Excessive smearing may suggest DNA fragmentation or degradation during extraction.
DNA concentration: The concentration of the extracted DNA can be determined using a spectrophotometer or fluorometric method. DNA yield may vary depending on the plant species, tissue type, sample condition, and efficiency of the extraction procedure.
DNA purity: Spectrophotometric analysis can be used to assess DNA purity through absorbance ratios. Under optimized extraction conditions, DNA preparations generally show acceptable A260/A280 and A260/A230 ratios, although expected values can vary depending on the sample and extraction method.
Overall result: A successful CTAB extraction should produce sufficiently concentrated, relatively pure, and intact genomic DNA that is suitable for downstream applications such as PCR, restriction digestion, sequencing, genotyping, and other molecular analyses.
Modifications of DNA Isolation from Plant Tissues
Several modifications of the conventional CTAB method have been developed to accommodate differences in plant tissue type, metabolite composition, sample preservation, and DNA quality. These modifications can improve DNA yield, purity, and compatibility with downstream molecular applications.
Increased NaCl concentration: Increasing the concentration of NaCl in the CTAB extraction buffer can improve the separation of DNA from polysaccharides by reducing their co-precipitation during alcohol precipitation. This modification is particularly useful for mucilaginous and carbohydrate-rich plant tissues, which often produce polysaccharide-contaminated DNA preparations (Sahu et al., 2016).
Addition of PVP (Polyvinylpyrrolidone): PVP can bind oxidized polyphenolic compounds released during tissue disruption. By reducing the interaction of these compounds with DNA, PVP can improve DNA purity, integrity, and amplifiability, particularly when extracting DNA from plants with high phenolic content (Sahu et al., 2016).
Extended lysis time: Increasing the incubation period in CTAB extraction buffer can enhance the disruption of rigid or difficult-to-lyse plant tissues and improve DNA release. This modification can be particularly useful for silica-dried, aged, or herbarium specimens, where cellular structures may have undergone extensive changes during preservation (Schenk et al., 2023a).
Modified CTAB buffer composition: The concentrations of CTAB, NaCl, EDTA, and other buffer components can be adjusted according to the characteristics of the plant material. Such adaptations, commonly described as modified CTAB protocols, are used to optimize DNA recovery and purity for particular plant taxa or challenging sample types (Schenk et al., 2023b).
Reduced tissue input: CTAB protocols can be optimized to obtain usable genomic DNA from small quantities of plant material. This is particularly valuable when working with rare species, limited specimens, valuable herbarium material, or minimally destructive sampling approaches (Doyle & Doyle, 1987).
Overall, these modifications allow the CTAB method to be adapted to different plant species and sample conditions, helping overcome common challenges associated with polysaccharides, polyphenols, rigid tissues, and limited starting material while maintaining DNA quality for downstream molecular analysis.
Troubleshooting of DNA Isolation from Plant Tissues
Low DNA yield
Likely causes: Incomplete cell lysis, insufficient tissue disruption, inadequate extraction conditions, or the use of old/degraded plant tissue.
Solutions: Grind the tissue thoroughly to obtain a fine powder, ensure complete mixing with CTAB buffer, and increase the lysis temperature or incubation time when appropriate.
DNA degradation
Likely cause: DNase activity during tissue processing or DNA extraction.
Solutions: Ensure that the extraction buffer contains sufficient EDTA, process samples promptly, minimize unnecessary handling, and avoid prolonged exposure of the extracted DNA to conditions that promote degradation.
Brown or highly viscous DNA preparation
Likely cause: Contamination with polyphenolic compounds and polysaccharides.
Solutions: Use freshly added β-mercaptoethanol to limit phenolic oxidation and consider incorporating PVP (polyvinylpyrrolidone) into the extraction protocol for phenolic-rich plant tissues. Additional organic extraction may also improve purity.
Poor PCR amplification
Likely cause: Carryover of salts, polysaccharides, phenolic compounds, CTAB, or other extraction contaminants that inhibit DNA polymerase activity.
Solutions: Perform an additional 70% ethanol wash, ensure complete removal of the wash solution, and verify DNA purity before PCR. If necessary, further purify the DNA before amplification.
RNA contamination
Likely cause: Insufficient removal of RNA during DNA extraction.
Solution: Treat the aqueous DNA-containing fraction with RNase A under appropriate conditions before DNA precipitation or during the purification workflow.
Overall troubleshooting principle: Successful plant DNA isolation depends on efficient tissue disruption, adequate removal of polysaccharides and phenolic compounds, protection against nuclease activity, and thorough removal of salts and other inhibitors. The appropriate modification should be selected according to the plant species and tissue characteristics (Doyle & Doyle, 1987; Sahu et al., 2016; Zymo Research, 2022).
Quality Assessment of the Isolated DNA
Agarose gel electrophoresis: Agarose gel electrophoresis is used to assess the integrity and degradation status of extracted genomic DNA. High-quality, intact DNA generally appears as a high-molecular-weight band near the well with minimal smearing, whereas extensive smearing may indicate DNA fragmentation or degradation.
Spectrophotometric analysis: DNA purity can be evaluated by measuring absorbance at 260 and 280 nm. An A260/A280 ratio of approximately 1.8 is generally considered indicative of relatively pure DNA, although the ratio should be interpreted alongside other quality measurements and the requirements of the downstream application (Biotech Beacon, 2025).
DNA concentration: Spectrophotometric or fluorometric measurements can also be used to determine the DNA concentration, helping establish whether sufficient DNA is available for subsequent molecular applications.
A260/A230 ratio: The A260/A230 ratio provides additional information about contamination by compounds such as salts, phenolic compounds, carbohydrates, and residual extraction reagents. A reduced ratio may indicate the presence of such contaminants.
Amplifiability: Successful amplification by PCR provides functional evidence that the extracted DNA is sufficiently pure and intact for enzymatic reactions. Poor or failed amplification may indicate DNA degradation or the presence of PCR-inhibitory contaminants, even when spectrophotometric measurements appear acceptable.
Overall assessment: Reliable evaluation of plant genomic DNA should ideally combine DNA concentration, purity measurements, agarose gel electrophoresis, and downstream PCR performance rather than relying on a single quality parameter.
Safety Tips and Precautions of DNA Isolation from Plant Tissues
Handle β-mercaptoethanol in a fume hood: β-Mercaptoethanol is a volatile, toxic, and strong-smelling reducing agent. Handle it in a properly functioning chemical fume hood while wearing appropriate personal protective equipment (PPE).
Wear appropriate personal protective equipment (PPE): Use laboratory gloves, a lab coat, and chemical-resistant eye protection to minimize exposure to extraction reagents and biological materials. Replace contaminated gloves immediately.
Handle chloroform carefully: Chloroform is a hazardous volatile organic solvent and should be handled in a chemical fume hood. Avoid inhalation and direct skin or eye contact, and keep containers closed when not in use.
Dispose of chloroform and organic solvent waste properly: Collect chloroform-containing waste in an appropriate designated organic-solvent waste container. Do not pour chloroform or other organic solvents down the sink. Follow your institution's chemical waste disposal procedures (Zymo Research, 2022).
Handle liquid nitrogen with appropriate protection: Use cryogenic gloves and suitable eye/face protection when handling liquid nitrogen. Avoid skin contact and work in a well-ventilated area because evaporating nitrogen can displace oxygen.
Exercise caution with heated CTAB buffer: When incubating CTAB extraction mixtures at elevated temperatures, use appropriate tube-handling equipment and avoid opening hot tubes abruptly to minimize the risk of splashes or exposure.
Prevent cross-contamination: Use clean or sterile equipment, properly labeled tubes, and fresh reagents where required. Keep samples and reagents organized to prevent accidental mixing or contamination.
Follow institutional laboratory safety procedures: Consult the relevant Safety Data Sheets (SDSs) for CTAB, β-mercaptoethanol, chloroform, isoamyl alcohol, and other reagents before use, and follow your laboratory's chemical, biological, and waste-management requirements.
Storage and Long‑Term Stability of Isolated DNA
Short-term storage at 4°C: Purified genomic DNA can generally be stored at 4°C for short-term use, particularly when the DNA will be used within a relatively short period. Repeated temperature fluctuations should be minimized.
Long-term storage at −20°C or −80°C: For extended storage, DNA is typically maintained at −20°C or −80°C to reduce degradation and preserve DNA integrity. −80°C storage may be preferred for long-term preservation of valuable samples.
Use TE buffer for DNA stability: Resuspending DNA in TE buffer (Tris-EDTA) can improve long-term stability. Tris helps maintain a suitable pH, while EDTA chelates divalent metal ions required by many nucleases, thereby helping protect DNA from enzymatic degradation.
Avoid repeated freeze–thaw cycles: Repeated freezing and thawing can contribute to DNA damage and should be minimized. For frequently used samples, preparing small aliquots can reduce the need to repeatedly thaw the entire DNA preparation.
Use nuclease-free water when appropriate: DNA may also be stored in nuclease-free water, particularly when downstream applications may be sensitive to EDTA. However, TE buffer generally provides greater protection against nuclease-mediated degradation.
Maintain proper labeling and documentation: Store DNA in clearly labeled, securely sealed tubes and record relevant information such as sample identity, concentration, extraction date, and storage conditions to maintain sample traceability.
Applications of DNA Isolation from Plant Tissues
PCR and qPCR: Isolated plant DNA provides a template for amplifying and detecting specific genetic sequences. PCR and qPCR can be used for gene detection, transgene verification, pathogen identification, molecular diagnostics, and functional genomics. qPCR can also be used to quantify target nucleic acid sequences, depending on the experimental design.
Genotyping and Phylogenetics: Plant genomic DNA is used for genotyping, population genetic studies, and genetic diversity analysis. Molecular markers derived from isolated DNA can support species and cultivar identification, phylogenetic tree construction, and investigation of evolutionary relationships among closely related plant taxa.
Plant Breeding and Biotechnology: DNA isolation supports molecular approaches in plant breeding, including marker-assisted selection, identification of desirable genetic traits, verification of genetically modified plants, trait introgression, and genetic characterization of breeding materials. These approaches contribute to the development of crop varieties with desirable characteristics such as improved yield and stress tolerance.
Herbarium and Conservation Genetics: DNA extracted from herbarium specimens and preserved plant material can be used to investigate historical genetic variation, identify species, study phylogeographic patterns, and support conservation genetics. Specialized extraction approaches can make molecular analysis possible even when only small quantities of degraded or historically preserved plant material are available (Schenk et al., 2023a).
Overall significance: Plant DNA isolation provides the molecular foundation for applications spanning genetic analysis, plant breeding, biodiversity research, taxonomy, conservation biology, biotechnology, and molecular diagnostics.
Advantages and Limitations of DNA Isolation from Plant Tissues
Advantages
Cost-effective: The CTAB method generally uses inexpensive and readily available laboratory reagents, making it suitable for routine plant DNA extraction, particularly in laboratories with limited resources.
Highly versatile: The method can be adapted to a wide range of plant species and tissue types, including fresh, dried, preserved, and metabolite-rich samples. Its buffer composition can also be modified according to the characteristics of the plant material.
Good DNA yield and purity: When appropriately optimized, CTAB extraction can produce high-quality genomic DNA by effectively removing common plant contaminants such as polysaccharides, proteins, and phenolic compounds.
Limitations
Labor-intensive: The conventional CTAB procedure involves several manual steps, including tissue grinding, incubation, organic extraction, centrifugation, DNA precipitation, washing, and resuspension. This increases hands-on laboratory time.
Use of hazardous chemicals: The protocol commonly involves chloroform and β-mercaptoethanol, which require appropriate handling, personal protective equipment, fume-hood use where applicable, and proper chemical-waste disposal.
Time-consuming: Multiple extraction, centrifugation, incubation, and precipitation steps make the conventional CTAB method relatively slow compared with some commercial or automated DNA extraction systems.
Limited suitability for high-throughput workflows: Because of its manual handling requirements and use of organic solvents, the conventional CTAB protocol can be less convenient for large-scale or highly automated DNA extraction workflows (Zymo Research, 2022).
Conclusion
The CTAB method remains a widely established technique for isolating genomic DNA from plant tissues because of its robustness, versatility, cost-effectiveness, and ability to address plant-specific extraction challenges.
Since its introduction by Doyle and Doyle (1987), the method has undergone numerous modifications to improve DNA yield, purity, and integrity, particularly when working with challenging samples containing high levels of polysaccharides, polyphenols, secondary metabolites, or degraded material.
Although CTAB extraction involves multiple manual steps and requires careful handling of certain chemicals, it continues to be widely used in academic, research, and molecular biology laboratories because it can be readily adapted to different plant species and sample types.
The continued use of CTAB-based extraction demonstrates the importance of understanding both the biochemical principles and practical considerations involved in plant DNA isolation. Proper optimization of tissue disruption, lysis, contaminant removal, precipitation, washing, and DNA storage is essential for obtaining DNA suitable for reliable downstream molecular applications.
References
Biotech Beacon. (2025). DNA isolation from plant tissue using CTAB method: Complete guide. https://www.biotechbeacon.com/dna-isolation-from-plant-tissue-using-ctab-method-complete-guide
Doyle, J. J., & Doyle, J. L. (1987). A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin, 19, 11–15.
Sahu, S. K., Thangaraj, M., & Kathiresan, K. (2012). DNA extraction protocol for plants with high levels of secondary metabolites and polysaccharides: A CTAB-based method. ISRN Molecular Biology, 2012, Article 205049. https://doi.org/10.5402/2012/205049
Schenk, J. J., Becklund, L. E., Carey, S. J., & Fabre, P. P. (2023). What is the “modified” CTAB protocol? Characterizing modifications to the CTAB DNA extraction protocol. Applications in Plant Sciences, 11(3), e11541. https://doi.org/10.1002/aps3.11541
Schenk, M. F., Thoen, M. A., Brøndum, R. F., Boyd, R. K., Suda, R., & Viljoen, C. (2023a). Optimizing the lysis step in CTAB DNA extractions of silica-dried and herbarium leaf tissues. Applications in Plant Sciences, 11(3), e12100. https://doi.org/10.1002/aps3.12100
Schenk, M. F., Thoen, M. A., Brøndum, R. F., Boyd, R. K., Suda, R., & Viljoen, C. (2023b). What is the “modified” CTAB protocol? Characterizing modifications to a classic DNA extraction method. Applications in Plant Sciences, 11(6), e12121. https://doi.org/10.1002/aps3.12121
Zymo Research. (2022). CTAB protocol for isolating DNA from plant tissues. https://www.zymoresearch.com/blogs/blog/ctab-protocol-for-isolating-dna-from-plant-tissues
Kathmandu University. (2025). Phenol–chloroform extraction manual for human urine samples. Unpublished laboratory protocol.
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