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Carbotrace are optotracers for anatomical mapping of carbohydrate structures in plants and for non-destructive composition analysis of glucans in bio-based materials and biofuels.

The optotracing technology entails the use of structure-responsive fluorescent molecules (optotracers) which become fluorescent when binding to a target. The excitation and emission spectra of the optotracer molecule contain information on the structure of their binding partner and their environment. This spectral fingerprint can be used to identify biomolecules in various types of materials.

Carbotrace is available in five variants which bind to repetitive motifs in proteins and carbohydrates. It has been shown that Carbotrace 680 binds to homoglucans and heteroglucans with a limited degree of branching. By using the spectral fingerprint from the excitation- and emission spectra, it was shown that the optotracer can differentiate between different types of glycosidic bonds. Using Multi-laser/Multi-detector imaging, non-destructive composition analysis of biomass has been performed such as differentiation of cellulose cell walls and starch granules in fresh potato as well as cellulose and lignin in in various types of biomass. Using fluorescence lifetime imaging in combination with Carbotrace 680, the mechanism of paper ageing was analysed. Further, Carbotrace 680 was used to visualise cellulose content in terrestrial plants (maize root and ulvophyceae) as well as algae and to characterise a cellulose immobilisation matrix in a microfluidic device. Carbotrace 480 was used as a tool for cellulose visualisation and to optimise cellulose extraction from macroalgae.

Carbotrace variant work in a wide range of salt and pH conditions. When the pH is altered during the experiment, pH controls should be included. Carotrace can be used with fluorescence plate readers, fluorescence microscopes and confocal laser scanning microscopes and fluorescence life time imaging.

Store your Carbotracer product in the fridge and use the opened container within 12 months. Carbotrace is for research use only and is not for resale.

Carbotrace Testing Service

Carbotrace Testing Service

Are your interested in composition analysis using Carbotrace, but lack necessary equipment or expertise?

With our Carbotrace Testing Service, we can help you get started!

If you send us your material, we will be able to label your sample with a Carbotrace variant of your choice and acquire data using our protocols and in-house equipment. As a result, you will receive a report including some processed data and all the raw data. Samples are able to be sent back or destroyed upon request when the project is completed.

Please click on the button to fill out an inquiry form.

If you fill out all fields, we will be able to get back to you with a detailed quote for a Carbotrace Testing Service tailored to your needs. If you only fill out your name and email, we will be happy to get in touch to discuss the best way forward.

Fill out Carbotrace Testing Service Request
Carbotrace Mix&Try

Carbotrace Mix&Try is our recommended option for starting out with using Carbotrace. It contains 10 µL of each variant. Testing each variant will allow you to determine which variant is best suited for your experiments and available instruments.

All Carbotrace variants label repetitive motifs in proteins and carbohydrates. Generally, Carbotrace labeling is strongest with protein aggregates. Homoglucans like cellulose are labeled reliably, albeit with lower quantum yield. Heteroglucans with low degree of branching like starches are labeled and can be distingiushed by their spectral fingerpint. Labelling of hemicelluloses is weak or absent.

Carbotrace 480 has been used to optimize a sustainable extraction method for cellulose hydrogels derived from microalgae and Carbotrace 680 has been used to image cellulose and starch components in plant cells and differentiate between those components using multi-laser/multi-detector analysis with confocal microscopy. Using a fluorescence spectrophotometer to acquire excitation and emission spectra, a unique spectral signature allows to differentiate between α(1-3), α(1-4), α(1-6), β(1-3), β(1-4) and β(1-6) linked glucans. As such, Carbotrace 680 labels cell wall components such as Cellulose, Chitin, Laminarin and the hemicellulose xyloglucan. Further, Carbotrace 680 has been used to determine relative carbohydrate content and elasticity of cell walls in plant material. Contact us to learn more about applications for Carbotrace.

Table: Excitation- and emission wavelengths as well as recommended filter sets.
Exmax Emmax Recommended filter-sets
Carbotrace 480 420 nm 480 nm DAPI
Carbotrace 520 460 nm 520 nm FITC, GFP
Carbotrace 540 480 nm 540 nm FITC, GFP, YFP
Carbotrace 630 520 nm 630 nm PI, Cy3, TxRed, mCherry, Cy3.5
Carbotrace 680 530 nm 680 nm PI, mCherry, Cy3.5
contact us for custom options.
Carbotrace 680

Carbotrace 680 is our red optotracer for labeling repetitive motifs in proteins and carbohydrates. Generally, Carbotrace labeling is strongest with protein aggregates. Homoglucans like cellulose are labeled reliably, albeit with lower quantum yield. Heteroglucans with low degree of branching like starches are labeled and can be distingiushed by their spectral fingerpint. Generally labelling of hemicelluloses is weak or absent.

Specifically, Carbotrace 680 has been used to image cellulose and starch components in plant cells and differentiate between those components using multi-laser/multi-detector analysis with confocal microscopy. Using a fluorescence spectrophotometer to acquire excitation and emission spectra, a unique spectral signature allows to differentiate between α(1-3), α(1-4), α(1-6), β(1-3), β(1-4) and β(1-6) linked glucans. As such, Carbotrace 680 labels cell wall components such as Cellulose, Chitin, Laminarin and the hemicellulose xyloglucan. Further, Carbotrace 680 has been used to determine relative carbohydrate content and elasticity of cell walls in plant material. Contact us to learn more about applications for Carbotrace 680.

Table: Excitation- and emission wavelengths as well as recommended filter sets.
Exmax Emmax Recommended filter-sets
Carbotrace 680 530 nm 680 nm PI, mCherry, Cy3.5

Carbotrace 680 is available in four different formulations (See volumes and prices in the drop-down list below): 

  • Aqueous: 1 mg/ml solution in ultrapure water. The product should be diluted 1:1000 before use. To prevent evaporation of the aqueous solvent, close the container carefully after use, spin down liquid and use up small volumes quickly.
  • DMSO: 1 mg/ml solution in DMSO to prevent solvent evaporation. The product should be diluted 1:1000 before use.
  • Solid: 1 mg solid lyophilised in a sterile injection bottle.
  • Drop&Shine:  5 ml ready-to-use product in mounting medium. Ideal for use in sections. Add a some Drop&Shine and mount your slide to analyse biomass or other cellulosic materials within minutes.
contact us for custom options.
Carbotrace 520

Carbotrace 520 is our green optotracer for labeling repetitive motifs in proteins and carbohydrates. Generally, Carbotrace labeling is strongest with protein aggregates. Homoglucans like cellulose are labeled reliably, albeit with lower quantum yield. Heteroglucans with low degree of branching like starches are labeled and can be distingiushed by their spectral fingerpint. Generally labelling of hemicelluloses is weak or absent. Contact us to learn more about applications for Carbotrace 520.

Table: Excitation- and emission wavelengths as well as recommended filter sets.
Exmax Emmax Recommended filter-sets
Carbotrace 520 460 nm 520 nm FITC, GFP

Carbotrace 520 is available as 1 mg/ml solution in ultrapure water (Aqueous) with volumes ranging from 10 - 200 µL (See volumes and prices in the drop-down list below). The product should be diluted 1:1000 before use. To prevent evaporation of the aqueous solvent, close the container carefully after use, spin down liquid and use up small volumes quickly.

contact us for custom options.
Carbotrace 630

Carbotrace 630 is our orange optotracer for labeling repetitive motifs in proteins and carbohydrates. Generally, Carbotrace labeling is strongest with protein aggregates. Homoglucans like cellulose are labeled reliably, albeit with lower quantum yield. Heteroglucans with low degree of branching like starches are labeled and can be distingiushed by their spectral fingerpint. Generally labelling of hemicelluloses is weak or absent. Contact us to learn more about applications for Carbotrace 630.

Table: Excitation- and emission wavelengths as well as recommended filter sets.
Exmax Emmax Recommended filter-sets
Carbotrace 630 520 nm 630 nm PI, Cy3, TxRed, mCherry, Cy3.5

Carbotrace 630 is available as 1 mg/ml solution in ultrapure water (Aqueous) with volumes ranging from 10 - 200 µL (See volumes and prices in the drop-down list below). The product should be diluted 1:1000 before use. To prevent evaporation of the aqueous solvent, close the container carefully after use, spin down liquid and use up small volumes quickly.

contact us for custom options.
Carbotrace 540

Carbotrace 540 is our yellow optotracer for labeling repetitive motifs in proteins and carbohydrates. Generally, Carbotrace labeling is strongest with protein aggregates. Homoglucans like cellulose are labeled reliably, albeit with lower quantum yield. Heteroglucans with low degree of branching like starches are labeled and can be distingiushed by their spectral fingerpint. Generally labelling of hemicelluloses is weak or absent.

Specifically, Carbotrace 540 has been used to determine the purity of cellulose extracted from macroalgae by allowing analysis of extracted carbohydrates in their polymeric state. In addition, Carbotrace 540 has shown to bind to the hemicellulose xyloglucan. Contact us to learn more about applications for Carbotrace 540.

Table: Excitation- and emission wavelengths as well as recommended filter sets.
Exmax Emmax Recommended filter-sets
Carbotrace 540 480 nm 540 nm FITC, GFP, YFP

Carbotrace 540 is available as 1 mg/ml solution in ultrapure water (Aqueous) with volumes ranging from 10 - 200 µL (See volumes and prices in the drop-down list below). The product should be diluted 1:1000 before use. To prevent evaporation of the aqueous solvent, close the container carefully after use, spin down liquid and use up small volumes quickly.

contact us for custom options.
Carbotrace 480

Carbotrace 480 is our blue optotracer for labeling repetitive motifs in proteins and carbohydrates. Generally, Carbotrace labeling is strongest with protein aggregates. Homoglucans like cellulose are labeled reliably, albeit with lower quantum yield. Heteroglucans with low degree of branching like starches are labeled and can be distingiushed by their spectral fingerpint. Generally labelling of hemicelluloses is weak or absent.

Specifically, Carbotrace 480 has been used to optimize a sustainable extraction method for cellulose hydrogels derived from microalgae. Contact us to learn more about applications for Carbotrace 480.

Table: Excitation- and emission wavelengths as well as recommended filter sets.
Exmax Emmax Recommended filter-sets
Carbotrace 480 420 nm 480 nm DAPI

Carbotrace 480 is available as 1 mg/ml solution in ultrapure water (Aqueous) with volumes ranging from 10 - 200 µL (See volumes and prices in the drop-down list below). The product should be diluted 1:1000 before use. To prevent evaporation of the aqueous solvent, close the container carefully after use, spin down liquid and use up small volumes quickly.

contact us for custom options.

Protocol I: Anatomical mapping of cellulose in plant tissues using Carbotrace 680

This protocol describes a non-destructive method of anatomical mapping of cellulose structures in plant tissues. Solutions and Reagents: Carbotrace 680 is provided as concentrated solution. The following common reagents are required (not supplied): Phosphate buffered saline (PBS), pH 7.4 Assay Procedure: Prepare thin sections of plant tissue. Dilute Carbotrace 680...
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Protocol II: Measurement of cellulose content in a sample using Carbotrace 680

This protocol describes how to use Carbotrace 680 to determine cellulose content in a sample. Solutions and Reagents: Carbotrace 680 is provided as concentrated solution. The following common reagents are required (not supplied): Phosphate buffered saline (PBS), pH 7.4 96-well plate (round bottom) Spectrophotometer Assay Procedure: Prepare a dilution series...
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A sustainable solution for transforming food waste into high-value hydrogels

A sustainable solution for transforming food waste into high-value hydrogels
Researchers from the University of York focused on creating hydrogels from blackcurrant pomace, a residue from the food supply chain. They aimed to explore sustainable methods for producing valuable biopolymers like cellulose through microwave-assisted hydrothermal fractionation, thus reducing waste. The team first isolated pectin at various temperatures, then used a...
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Carbotrace for differentiation between lignin and cellulose in renewable resources

Carbotrace for differentiation between lignin and cellulose in renewable resources
A study by researchers from KTH explores Lupinus angustifolius (a type of lupin) as a sustainable source of lignocellulose materials. In order to reduce reliance on forest wood and utilise agricultural residues to produce microfibrillated cellulose (MFC), the team analysed the anatomical structure and chemical composition of the plant, classifying...
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Carbotrace as a tool for the study of cellulose oxidation and paper aging

Carbotrace as a tool for the study of cellulose oxidation and paper aging
Since its invention, paper has skyrocketed technological advances as a key medium for recording thoughts and ideas. However, over time paper ages and oxidation leads to loss of strength and renders sheets of paper brittle. Therefore, the study of paper aging becomes relevant not only for the preservation of historical...
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Making greener cellulose hydrogel

Making greener cellulose hydrogel
The use of Microalgae as a renewable resource has attracted large interest due to their potential for use in the renewable energy, biopharmaceutical, and nutraceutical industries. Microalgae are renewable, sustainable, and economical sources of biofuels, bioactive medicinal products, and food ingredients. Recently, researchers around Professor Avtar Singh Matharu from the...
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Carbohydrate content and elasticity of cell walls in elongating maize root

Carbohydrate content and elasticity of cell walls in elongating maize root
Investigation of plant growth and development is now more than ever an important field of study following the many challenges faced in agriculture. Accurate methods and computational models of plant cell- and organ change during growth are therefore essential to tackle problems like increased food demand in the wake of...
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Breaking barriers in early detection of cancer

Breaking barriers in early detection of cancer
The ability to detect and isolate circulating tumor cells from blood could be a life-saving diagnostic tool for early detection of cancer and downstream analysis of tumor growth and metastasis. Researchers from the lab of Aman Russom at the Royal Institute of Technology (KTH) in Sweden have engineered a microfluidic...
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Carbotrace 540 was used to determine purity of cellulose extracted from sea lettuce

Carbotrace 540 was used to determine purity of cellulose extracted from sea lettuce
The video clip shows the eco-friendly extraction of of pure cellulose from green macroalgae from the Swedish west coast. Researchers from the Royal Institute of Technology and Karolinska Institutet in Sweden reported about an environmentally friendly process to obtain pure cellulose nanofibrils from the green macroalgae ulva lactuca. Cellulose was...
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Carbotrace-like molecules allow highly detailed visualization of cellulose in plant cells

Carbotrace-like molecules allow highly detailed visualization of cellulose in plant cells
If you always wondered why it is so difficult to switch to plants for producing renewable resources and how Carbotrace can help in the process, the video from Ben@SwedishMedicalNanoscienceCenter explains: "The complicated thing about plants and biofuels." A study from 2018 published in Nature Scientific Reports used a Carbotrace-like molecule...
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Carbotrace 680 for carbohydrate anatomical mapping

Carbotrace 680 for carbohydrate anatomical mapping
The video clip below illustrates why our current model of Take, Make, Dispose is draining our planet of natural resources and why circular economy and renewable resources are the answer. Learn how Carbotrace can help to identify renewable resources in plant biomass and support circular economy in the food &...
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Monitoring heat-induced swelling of Starch granules

Monitoring heat-induced swelling of Starch granules
As starch is a thermo-sensitive material that undergoes drastic morphological changes when heated, it is even possible to monitor starch reorganization during heat-induced swelling. When freshly extracted potato starch granules marked with Carbotrace 680 are heated from 20 °C to 90 °C and imaged with a confocal microscope, it becomes...
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Carbotrace fluorescence spectra

Carbotrace fluorescence spectra
We named our Carbotrace molecules after their peak emission wavelength when they are bound to their target. That means, when Carbotrace is bound to a target, it will emit fluorescence at peak emission indicated by the number associated with its name. To view the excitation and emission spectra, please select...
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Carbotrace 680 for monitoring starch

Carbotrace 680 for monitoring starch
Detection of polysaccharides using Carbotrace 680 relies on regularly occurring units joined by glycosidic linkages. The specific type of glycosidic linkage and the occurrence of branches allow Carbotrace 680 to differentiate between different types of glucans. By means of its structure-responsive properties, Carbotrace 680 can differentiate between β(1-4) linked cellulose...
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Testimonial - Reina Tanaka

Reina Tanaka about Carbotrace & Ebba Biotech Research service: “Ebba Biotech helped us by performing analyses on Tunicate derived Cellulose nanocrystals and softwood TEMPO Oxidised Cellulose nanofibrils to determine the best Carbotrace molecule for our analysis. This encompassed both spectrophotometric testing and data analysis. This service was instrumental for us...
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Testimonial - Frederik Zitzmann

Prof. Avtar Matharu & Frederik Zitzmann (doctoral student) about Carbotrace 480: ”We used Carbotrace 480 as an analytical tool to map cellulose content and distribution in our defibrillated cellulose samples which we derived from microalgae. Due to the complex structure of the algal cell wall we had major difficulties in...
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Testimonial - Saga Jakobsson

Saga Grånäs Jakobsson about Carbotrace 680: “I used Carbotrace 680 in my master thesis project where I investigated hemicelluloses and nanocellulose. I was able to detect hemicelluloses xyloglucan and galactomannan with Carbotrace 680. Also, I was able to use Carbotrace 680 to assess cellulose presence in commercially available samples and...
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Testimonial - Aristoteles Goes-Neto

Prof. Aristóteles Góes-Neto & Thairine Mendes-Pereira (PhD student) about Carbotrace 680: "We used Carbotrace 680 to label fungal cell wall chitin and spider exoskeleton chitin. More specifically, we studied the host-pathogen interaction of the pathogenic fungal genus Gibellula (Ascomycota) and the Pholcidae family of spiders. In our hands, Carbotrace 680...
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Testimonial - Ulrica Edlund

Prof. Ulrica Edlund about Carbotrace 540: "We were studying the biochemical composition of Ulva lactuca (a.k.a. Ulva fenestrata), which is still not fully known. Our hypothesis, and we wanted to prove, was the presence of cellulose and we knew that optotracing was probably the best method for analysis to distinguish...
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Testimonial - Ferdinand X. Choong

Dr. Ferdinand X. Choong about Carbotrace 680: "It really made carbohydrate detection so much easier! Thinking back on how to detect starch you had to use iodine solutions - that was kind of messy - and now Carbotrace 680 just works - simple and accurate." Dr. Ferdinand X. Choong is...
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Testimonial - Ulrica Edlund

Prof. Ulrica Edlund about Optotracing: Ulrica Edlund is Professor in Polymer technology at KTH Royal Institue of Technology and Vice Director at the Center for the Advancement of Integrated Medical and Engineering Sciences (AIMES) at Karolinska Institutet in Stockholm, Sweden. Optotracing in biorefinement of Canola Straw. Thin sections of canola...
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Testimonial - Liudmila Kozlova

Dr. Liudmila Kozlova about Carbotrace 680: "We have used Carbotrace 680 to resolve the cellular structure of studied plants. It is possible due to the high content of cellulose in plant cell walls. Moreover, the fluorescence intensity of Carbotrace 680 can be used for the determination of cell wall thickness...
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Testimonial - Tharagan Kumar

Tharagan Kumar about Carbotrace 680: Tharagan Kumar is doctoral student at KTH Royal Institute of Technology, Division of Nanobiotechnology located at SciLifeLab in Solna, Sweden. Tharagan sent us an image about how he was using Carbotrace 680 in his work. Optimization of an immobilization matrix made of cellulose nanofibrils in...
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Incorporating Carbotrace as a problem solver in a biorefinery design process

Ebba Biotech welcomes you to tune in to our webinar featuring Dr. Frederik Zitzmann from the University of York. During his talk titled "Incorporating Carbotrace as a problem solver in a biorefinery design process.", Dr. Zitzmann will discuss how we can use novel technologies such as visualising with Carbotrace to...
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Cellulose in Cancer Diagnostics

Ebba Biotech welcomes you to listen to our newest webinar featuring Dr. Tharagan Kumar presenting his work from his time at KTH, the Royal Institute of Technology in Stockholm Sweden. During this talk titled “Cellulose in Cancer Diagnostics”, Dr. Kumar will be presenting his work breaking barriers in diagnostics using...
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Spectral identification and recovery of polysaccharides from biorefining

Ebba Biotech welcomes you to listen to Professor Ulrica Edlund from the Royal Institute of Technology (KTH) in Stockholm and her revolutionary work with Carbotrace and nano-materials. The webinar titled "Selective method for spectral identification and recovery of polysaccharides from biorefining" explains how she uses Optotracers, like Carbotrace, in her...
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Optotracers - multifunctional fluorescent tracers

On the first of June 2021, Ferdinand Choong, Ebba Biotech's co-founder, and Assistant Professor at Karolinska Institutet and AIMES (Center for the Advancement of Integrated Medical and Engineering), presented his research using Ebba Biotech's optotracers at the digital event Lab & Diagnostics of the Future 2021, held by Life Science...
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Carbotrace 540 helps to determine purity of plant raw materials

Researchers from the Royal Institute of Technology and Karolinska Institutet in Sweden reported about an environmentally friendly process to obtain pure cellulose nanofibrils from the green macroalgae ulva lactuca (Wahlström et al. (2020) Cellulose 27, 3707–3725). Analysis of the monosugars revealed that the extracted cellulose contains mainly glucose, but also...
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Carbotrace-like molecules for visualization of cellulose in plants

A research paper (Choong et al. (2018) Scientific Reports, 8, 3108) uses a Carbotrace-like molecule to visualize the location and structure of cellulose in plant cells. The video below explains why it is so difficult to switch to plants for producing renewable resources and how Carbotrace can help in the...
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Carbotrace supports circular economy

In a research paper published in Cellulose (Choong et al. (2019) Cellulose, 26, 4253–4264), our structure-responsive optotracer molecule Carbotrace 680 was used to demonstrate the potential of optotracing for carbohydrate anatomical mapping and spectral imaging. The video clip below illustrates why our current model of Take, Make, Dispose is draining...
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We named our Carbotrace molecules after their peak emission wavelength when they are bound to their target. That means, when Carbotrace is bound to a target, it will emit fluorescence at peak emission indicated by the number associated with its name.

To view the excitation and emission spectra, please select your Carbotrace below :

Excitation (blue lines) and emission (red lines) spectra of unbound Carbotrace (dotted lines) and Carbotrace bound to a target (solid lines).

2024

  • Schmidt, A. E. M., Choong, F. X., Richter‐Dahlfors, A., & Edlund, U. (2024). Defibrillated Lignocellulose Recovery Guided by Plant Chemistry and Anatomy – A Pioneering Study with Lupinus angustifolius. Advanced Sustainable Systems. https://doi.org/10.1002/adsu.202300632
  • Ferrara, V., Vetri, V., Pignataro, B., Chillura Martino, D. F., & Sancataldo, G. (2024). Phasor-FLIM analysis of cellulose paper ageing mechanism with carbotrace 680 dye. International Journal of Biological Macromolecules , 260, 129452. https://doi.org/10.1016/j.ijbiomac.2024.129452

2023

  • Inthalaeng, N., Dugmore, T. I. J., & Matharu, A. S. (2023). Production of Hydrogels from Microwave-Assisted Hydrothermal Fractionation of Blackcurrant Pomace. Gels, 9(9). https://doi.org/10.3390/gels9090674
  • Petrova, A., Ageeva, M., & Kozlova, L. (2023). Root growth of monocotyledons and dicotyledons is limited by different tissues. Plant Journal, 116(5), 1462–1476. https://doi.org/10.1111/tpj.16440
  • Holzinger, A., Plag, N., Karsten, U., & Glaser, K. (2023). Terrestrial Trentepohlia sp. (Ulvophyceae) from alpine and coastal collection sites show strong desiccation tolerance and broad light and temperature adaptation. Protoplasma. https://doi.org/10.1007/s00709-023-01866-2

2022

  • Zitzmann, F. L., Ward, E., & Matharu, A. S. (2022). Use of Carbotrace 480 as a Probe for Cellulose and Hydrogel Formation from Defibrillated Microalgae. Gels, 8(6), 383. https://doi.org/10.3390/gels8060383
  • Petrova, A., Sibgatullina, G., Gorshkova, T., & Kozlova, L. (2022). Dynamics of cell wall polysaccharides during the elongation growth of rye primary roots. Planta, 255(5), 108. https://doi.org/10.1007/s00425-022-03887-2

2020

  • Petrova, A., Gorshkova, T., & Kozlova, L. (2020). Gradients of cell wall nano-mechanical properties along and across elongating primary roots of maize. Journal of Experimental Botany. https://doi.org/10.1093/jxb/eraa561
  • Kumar, T., Soares, R. R. G., Dholey, L. A., Ramachandraiah, H., Aval, N. A., Aljadi, Z., Pettersson, T., & Russom, A. (2020). Multi-layer assembly of cellulose nanofibrils in a microfluidic device for the selective capture and release of viable tumor cells from whole blood. Nanoscale, 42. https://doi.org/10.1039/d0nr05375a

2019

  • Choong, F. X., Lantz, L., Shirani, H., Schulz, A., Nilsson, K. P. R., Edlund, U., & Richter-Dahlfors, A. (2019). Stereochemical identification of glucans by a donor–acceptor–donor conjugated pentamer enables multi-carbohydrate anatomical mapping in plant tissues. Cellulose, 26(7), 4253–4264. https://doi.org/10.1007/s10570-019-02381-5