Skip to content
Frontiers
Search

Mini Review

Frontiers in Astronomy and Space Sciences ·

Volume 13 - 2026 | https://doi.org/10.3389/fspas.2026.1843790

Building biofilm: engineering for life off Earth

  • Katherine J. Baxter1*corresponding author
  1. School of Molecular Biosciences, College of Medical Veterinary and Life Sciences, University of Glasgow, Glasgow, United Kingdom
Article metrics0 viewsView details
PDFCiteMetrics

Abstract

Most microbial life on Earth exists as biofilms, large interkingdom microbial communities living within a self-synthesised matrix of extracellular polymeric substances. These living structures are critical to the survival of all life on Earth, driving biogeochemical processes within ecosystems and performing key functions in the maintenance of animal and plant health. As powerhouses of these life-enabling processes, biofilms are an untapped resource which can be utilised to address several challenges of human space exploration. To explore this concept, this mini review first presents the viewpoint of biofilms as a multicellular entity akin to rudimentary tissue and explains why they should be investigated through this lens. This is followed by an overview of current biofilm engineering technologies, the gaps in our understanding, and the highlighting of key research priorities. Finally, this review discusses how biofilm engineering can be used to reduce health risk, promote sustainable human spaceflight, and support exploration of other planetary environments.

Keywords

1 Introduction

Biofilms are the dominant form of microbial life on Earth (Penesyan et al., 2021) and are complex interkingdom multispecies microbial communities embedded in a self-synthesised matrix of extracellular polymeric substances (EPS) (Sadiq et al., 2022). Far from being simple microbial aggregates, biofilms underpin biogeochemical cycling, drive ecosystem resilience and play essential roles in the health of plants, animals and humans (Deveau et al., 2018). Yet despite their centrality to life on Earth, biofilms remain an underexploited resource in the context of human space exploration. Their inherent adaptability, metabolic breadth and capacity for self-organisation position them as uniquely powerful living technologies for sustaining life and enabling construction beyond Earth.

A growing body of evidence demonstrates that biofilms exhibit structural, functional and developmental features analogous to early multicellular tissues (Nikolaev and Plakunov, 2007; Aguilar et al., 2015; Espinoza Miranda et al., 2025; Penesyan et al., 2021). They generate spatially organised subpopulations, coordinate behaviour through chemical and mechanical cues, and construct extracellular matrices that provide mechanical stability, adhesion and information flow. These parallels suggest that biofilms can be productively reframed not as problematic contaminants or loose microbial assemblages, but as rudimentary tissues with emergent properties that arise from collective organisation. This conceptual shift provides a powerful foundation for engineering biofilms in ways that mirror tissue engineering, by manipulating biochemical, biophysical and spatial cues to guide community behaviour and matrix deposition (Bakhshandeh et al.,.2023). Figure 1 provides and overview of these tissue-like properties of biofilms, along with the multilevel engineering strategies available, and the major application areas where these systems could support space exploration.

Biofilms as rudimentary tissues and astro-pioneer systems. Created in BioRender. Baxter, K. (2026) [https://BioRender.com/n1goapk](https://BioRender.com/n1goapk).
Figure 1. Biofilms as rudimentary tissues and astro-pioneer systems. Created in BioRender. Baxter, K. (2026) https://BioRender.com/n1goapk.

At the same time, advances in synthetic biology, materials science and spatial patterning technologies have expanded our capacity to manipulate biofilms at the level of cells, matrix components and 3D architecture. However, these approaches remain largely modular, targeting isolated aspects of biofilm systems rather than the collective dynamics that define biofilm function. To unlock the full potential of biofilms as living technologies there is a pressing need to integrate these engineering strategies with a deeper mechanistic understanding of biofilm ecology, community interactions and environmental responsiveness. To this end, this mini review explores why biofilms should be viewed as multicellular entities, what our current biofilm engineering capacities and knowledge gaps are, and discusses how engineered biofilms could be pivotal in space exploration.

2 Why view biofilms as a multicellular entity?

The idea of biofilm as a form of multicellularity or rudimentary tissue is not new (Nikolaev and Plakunov, 2007; Penesyan et al., 2021). Indeed, beyond what has already been discussed in the literature, biofilms share tissue-like features somewhat akin to loose connective tissue, which provides mechanical stability and scaffolding throughout the body (Abbott et al., 2013). For example, collagen-like proteins present in fungi and bacteria are required for biofilm formation and host integrin binding (Soloviev et al., 2007; Zhao et al., 2015; Kurakado et al., 2018; Abdel-Nour et al., 2019), reflecting the role of collagen-integrin interactions in tissue growth and maintenance (Heino, 2014; Quan et al., 2025), and a variety of biofilms contain mucopolysaccharides similar to that of the viscous component of connective tissue (Yadav et al., 2013; Felz et al., 2020; Knott et al., 2021; Fede et al., 2021). At the basic level, both metazoan tissue and biofilms can be described as a collection of cells maintained within an extracellular matrix. In tissues, subpopulations of cells with specialised functions arise through differentiation driven by external cues and conserved signalling systems co-ordinated across spatiotemporal contexts (Doupé, and Perrimon, 2016; De Belly et al., 2022; Espina et al., 2023), tissue-specific architecture arises though interplay of these two contributing elements. In biofilm, an analogous combination of events occurs to establish biofilm architecture. Initial microbial populations giving rise to biofilms are predominantly multispecies (Sadiq et al., 2022), introducing an equivalent heterogeneity to early tissue differentiation. As biofilms mature, internal biofilm microenvironments driven by gradients of oxygen, nutrient, waste and signalling molecules (Pisithkul et al., 2019; Flemming et al., 2023; Jo et al., 2022) produce cellular subpopulations with differential gene expression. For example, in Pseudomonas aeruginosa, the RmcA-mediated redox system in hypoxic and anoxic biofilm microenvironments drives alterations in biofilm matrix production, resulting in changes to surface morphology and increased area-to-volume ratio (Rossi et al., 2023), and in both Bacillus subtilis and Clostridium difficile, spatial regulation of cell differentiation within the biofilm is induced by differential phosphorylation of the conserved master regulator Spo0A. In these species, exopolysaccharide biosynthesis genes are indirectly induced in cell populations with low to intermediate Spo0A phosphorylation levels, whereas the sporulation cascade is induced in those cells with high phosphorylation levels (Chen Z. et al., 2022; DiCandia et al., 2024). Another example is the universal bacterial second messenger c-di-GMP, which determines cell fate and metabolic activity within biofilms in response to nutrient limitation in range of bacterial species including C. difficile (Purcell et al., 2017), P. aeruginosa (Lichtenberg et al., 2022; Zhang et al., 2022) and Shewanella putrifaciens (Sun et al., 2025). Furthermore, in archaea, cell subpopulations arise in response to mechanical stress to produce tissue-like structures (Rados et al., 2025), and cellular differentiation within colony biofilms of the unicellular yeast Saccharomyces cerevisiae is orchestrated by Gcn4p (Váchová et al., 2024), an orthologue of c-Jun, a key regulator of tissue development in mammals (Meng and Xia, 2011).

Across bacteria, archaea and fungi, these differentiation mechanisms demonstrate that biofilms deploy regulatory logic typically associated with early multicellular development. Taken together, these examples illustrate how closely processes within biofilms parallel cell fate specification in developing tissues, where phosphorelay cascades and mechanosensing events determine cell fates (Gudipaty et al., 2017; Xue et al., 2025).

Beyond these structural and functional parallels, biofilms also exhibit higher-order behaviours that align them with early multicellular systems. Biofilms generate and interpret gradients of oxygen, nutrients, pH and signalling molecules which act in a morphogen-like way (Serra and Hengge, 2014; Jo et al., 2022), providing positional information that drive spatially distinct phenotypes in a manner analogous to developmental patterning in tissues (Simsek and Özbudak, 2022). Mechanical forces such as shear and compression feed back into matrix production and architectural remodelling (Yaman et al., 2019; Lipke, 2025; Karim and LaJeunesse, 2026; Wong et al., 2023) mirroring the mechanobiological responses seen in tissues (Espina et al., 2023; Mao and Wickström, 2024). Functionally, biofilms display clear division of labour, with matrix producing cells, metabolic specialists, motile dispersal subpopulations and persister cells (van Gestel et al., 2015; Flemming et al., 2023), similar to cellular specialisation within tissue. These behaviours are not incidental; biofilms represent the ancestral organisational state from which multicellularity evolved. Their extracellular matrices function as evolutionary precursors to the scaffolding, adhesion and signalling functions of metazoan extracellular matrix.

If we consider biofilm as a form of tissue, we can apply division of labour frameworks and tissue engineering approaches to develop novel technologies for the enablement of space exploration. The fundamental role of biofilms in the support of life on Earth and their ubiquity across ecological niches makes them ideal tools for this purpose. Depending on the mission need, synthetic biofilm communities can be built from the bottom-up to produce specific materials, undertake a task or be environmentally deployed to support spacecraft maintenance or habitat development.

3 Current biofilm engineering approaches

Our ability to manipulate biofilms currently sits across three structural levels: the cells themselves, the extracellular matrix and control of 3D spatial organisation. Each layer of engineering has seen substantial progress, but these advances remain largely modular. To reach whole biofilm engineering, the equivalent of tissue-level control, we need to integrate these layers rather than treat them as isolated engineering targets.

A broad range of microorganisms now function as chassis organisms for synthetic biology. These include Escherichia coli (Matsumoto et al., 2017), Bacillus subtilis (Cao et al., 2025); Corynebacterium glutamicum (Chai et al., 2021); Pseudomonas putida (Martinez Garcia and de Lorenzo, 2024); Vibrio natriegens (Liu et al., 2022); Halomonas bluephagenesis (Chen G. Q. et al., 2022) and Thermus thermophilus (Liang et al., 2025), with Saccharomyces species (Durmusoglu et al., 2023; Long et al., 2024) and Pichia pastoris (Xue et al., 2017; Vijaykumar and Venkataraman, 2024) as fungal options. These microorganisms are used to produce pharmaceuticals (Liu et al., 2022; Yang et al., 2024) chemicals and fuels (Gao et al., 2016; Della Valle et al., 2024; Azevedo et al., 2025) and plastic alternatives (González-Rojo et al., 2024). Crucially, some of these organisms are extremophiles. H. bluephagenesis and T. thermophilus tolerate high salt and elevated temperatures, respectively, with the former being adapted for production of amino acids and biodegradable polyesters at lower salt concentrations (Zhang et al., 2023). Across these chassis, several shared characteristics underpin their suitability for engineered biofilms. Fast-growing organisms such as V. natriegens and E. coli offer rapid design-build-test cycles and extensive genetic toolkits (Faber et al., 2025; Cooke et al., 2025), whereas B. subtilis, C. glutamicum and P. putida provide robust metabolic versatility and well-characterised regulatory elements (Cao et al., 2025; Chai et al., 2021; Martinez Garcia and de Lorenzo, 2024). Extremophiles such as H. bluephagenesis and T. thermophilus contribute valuable environmental tolerance, including high salinity or elevated temperature resistance (Chen G. Q. et al., 2022; Liang et al., 2025) while fungal systems such as Saccharomyces and Pichia offer eukaryotic secretion capacity and strong protein-production pipelines (Durmusoglu et al., 2023; Long et al., 2024; Xue et al., 2017; Vijaykumar and Venkataraman, 2024). Collectively, these chassis span a useful range of growth rates, genetic tractability, environmental resistance and metabolic breadth, providing a flexible foundation for engineering biofilms capable of operating in non-Terran environments. These key parameters should guide organism selection for space-relevant applications.

The extracellular polymeric substances (EPS) of biofilms are the structural and functional backbone of biofilms. Amyloid fibres and polysaccharides components confer structural integrity and surface interactions and have become major engineering targets. Curli, the archetypal Gram-negative amyloid, has been adapted for a striking range of functions. These include microplastic degradation (Zhu et al., 2022), covalent surface attachment and scaffolding (Nguyen et al., 2014), metal binding for corrosion protection (Li et al., 2024), heavy metal bioremediation (Tay et al., 2017; Kumari et al., 2025) and targeted drug delivery (Praveschotinunt et al., 2019). Parallel efforts in Gram-positive systems, Bap in Staphylococcus spp. (Matilla-Cuenca et al., 2022) and TasA in B. subtilis (Ren et al., 2025) are emerging but less mature. Beyond their role in biofilms, amyloids have been used directly as biodegradable plastics (Duraj-Thatte et al., 2021) and medical hydrogels (Wei et al. (2025). Polysaccharide engineering methods differ from those used for amyloid adaptation. Unlike amyloid protein display systems where enzymatic domain or surface affinity sequences are genetically incorporated into the protein, polysaccharide display systems rely on post-production processing to immobilise functional molecules (Dong et al., 2020). Other polysaccharide-focused engineering augments overall biofilm performance, such as increased functionality in fermentation processes (Pan et al., 2025) or water treatment processes (Kim et al., 2023). These approaches demonstrate that the matrix is a tuneable biochemical interface which can be utilised for a broad range of functionality.

At the level of global biofilm architecture, spatial control technologies push biofilms closer to engineered tissues by allowing us to dictate where cells attach, grow and interact. 3D printing and Biofilm lithography are the current methods available which allow the spatial distribution of cells within a matrix. 3D printing uses bioinks, which are either a hydrogel seeded with microorganisms (Johnston et al., 2020; Aliyazdi et al., 2023; Denton et al., 2024) or use the intrinsic biofilm forming capacity of the organism (Schmeiden et al., 2018; Huang et al., 2019). However, 3D printing using bioinks does not have the same degree of structural control as biofilm lithography, which uses optogenetics to drive biofilm deposition. In this method, molecular components of the planktonic to biofilm switch within the organism are engineered to be light-responsive. Light exposure triggers attachment and matrix production only in defined regions, enabling construction of monospecies and multispecies biofilm architectures (Jin and Riedel-Kruse, 2018; Jin and Riedel-Kruse, 2024). This is the closest analogue we currently have to tissue patterning. However, these technologies operate on isolated layers of the biofilm system rather than the collective behaviours that define biofilm function.

Despite these advances, we are still engineering components, not biofilms. Biofilms are emergent systems shaped by multispecies interactions, environmental gradients, and dynamic feedback loops. Variation into the system is introduced at both community and environmental level, making them inherently heterogeneous and hard to study as the same microbial community has the capacity to produce a different biofilm if environmental conditions change (Bjarnsholt et al., 2022). This heterogeneity underpins their behaviour as biofouling agents (Kochina et al., 2022) and infection risk (Percival et al., 2015; Maillard and Centeleghe, 2023) and is the primary barrier to deploying them as reliable technologies.

To transition from modular manipulation of biofilm components to true whole-biofilm engineering, several research priorities must be addressed. These priorities centre on resolving the fundamental knowledge gaps that currently prevent predictable, reproducible control of biofilm behaviour across environments, including non-Terran settings. Indeed, several reviews of space-flown biofilm experiments highlight the need for further study (Senatore et al., 2018; Prasad et al., 2021; Baxter et al., 2026). An eight-point priority list is detailed in Table 1, defining the research needed to transform biofilms from complex natural systems into predictable, programmable, and mission-ready living technologies for space exploration.

TABLE 1. Biofilm research priorities to enable space exploration.

Priority 1: High-resolution mapping of intracommunity interactions
Priority 2: Mechanistic understanding of environmental feedback loops
Priority 3: Gravity-dependent biofilm biology
Priority 4: Predictive modelling and control frameworks
Priority 5: Engineering tools for community-level manipulation
Priority 6: Standards, reproducibility, and open biofilm engineering platforms
Priority 7: Evolutionary stability and long-term functional reliability
Priority 8: Integration with spacecraft life-support, manufacturing and operational systems
Biofilms function through dense networks of cross-feeding, metabolic complementarity, antagonism, and cooperative signalling. These interactions remain poorly resolved, particularly in multispecies, interkingdom systems. Quantitative mapping of these networks using multi-modal omics and spatial transcriptomics is essential for identifying the regulatory nodes that can be engineered for community-level control
Biofilms integrate chemical, physical and mechanical cues into coordinated collective responses. Yet the mechanistic basis of these feedback loops and how gradients, shear forces, hydration dynamics and nutrient fluxes shape gene expression and matrix deposition remains incomplete. Decoding these relationships is critical for engineering biofilms that maintain function under fluctuating or extreme conditions, including microgravity, partial gravity, and radiation exposure
Biofilms behave differently along the gravity continuum, with alterations in matrix architecture and differential gene expression. Systematic investigation of how gravity shapes biofilm development, metabolism and resilience is required to design communities that remain stable and functional in spaceflight and planetary environments
Whole-biofilm engineering will require computational models capable of predicting emergent behaviours from cellular-level rules. Agent-based models, metabolic network reconstructions, and hybrid mechanistic-machine learning models are needed to integrate spatial, metabolic and regulatory data into actionable design frameworks. These models must be validated experimentally across environmental conditions relevant to space exploration
Current technologies target cells, matrix components, or spatial patterning in isolation. New tools are needed to modulate community composition, metabolic fluxes, matrix dynamics, and spatial organisation simultaneously. This includes synthetic gene circuits that operate across species boundaries, optogenetic or chemical control systems that function in 3D architectures, and chassis organisms engineered for interspecies communication and cooperative division of labour
Biofilm research suffers from variability in experimental conditions, strain selection, and environmental parameters. Establishing standardised protocols, reference communities and open-access datasets will be essential for reproducibility and for enabling comparison. These frameworks will accelerate the development of biofilm technologies suitable for mission-critical operations
Engineered biofilms must remain stable and predictable over extended timescales, yet the evolutionary dynamics of multispecies communities under microgravity, radiation and closed-loop conditions remain poorly understood. These uncertainties intersect with mission-critical risks, including pathogenicity, biofouling, corrosion and degradation of life-support infrastructure. Addressing this requires mechanistic insight into long-term evolutionary trajectories and the development of stabilisation and containment strategies, namely, genetic safeguards, ecological buffering and continuous monitoring systems to ensure safe and reliable operation in space habitats
The interfaces between engineered biofilms and spacecraft environmental control, water recovery, air revitalisation and in-situ manufacturing systems are not yet defined or validated. Without a deep functional understanding of how biofilm activity influences system-level parameters such as humidity, gas flux and fouling risk, integration remains a major barrier to deployment. Standardised interfaces, validated operational envelopes and hardware co-design frameworks are needed to ensure biofilm-based technologies can be safely incorporated, monitored and maintained within spaceflight infrastructure

4 Engineering biofilm for space exploration

Viewing biofilms as rudimentary tissue provides a powerful conceptual and practical framework for their use in space exploration. By treating biofilms as a form of multicellularity we can borrow directly from tissue engineering which routinely manipulates biochemical and biophysical cues to guide cell behaviour and matrix deposition (Bakhshandeh et al., 2023).

Applying tissue engineering principles to biofilms enables the creation of living construction materials that self-assemble, self-repair and adapt to environmental stressors. Emergent hybrid technologies of biofilm-building materials are in development (Răut et al., 2021; Ertelt et al., 2022; Li et al., 2022; Poohphajai et al., 2024; Wang et al., 2022) and engineered amyloid fibres and polysaccharides have already demonstrated the capacity for material augmentation (Zhu et al., 2022; Li et al., 2024). In space, these capabilities could be developed to produce composite materials with radiation-attenuating capacities, regolith-binding bio-cements, or flexible biopolymer coatings for habitat walls. Patterned distribution of biofilms onto structural substrates through future versions of Biofilm lithography and 3D printing can offer spatial control of these hybrid structural engineering approaches. These methods would allow for the development of living barrier systems., for example, layered biofilms could be used to bind and sequester molecules such as perchlorates to prevent corrosion or material degradation, and radiation shielding derived from already present resistance mechanisms (Ragon et al., 2011; Bratkic et al., 2024; Rahayu et al., 2025; Shrestha et al., 2025) could be augmented for the radiation profiles of specific non-Terran environments. As components of hybrid materials in spacecraft and habitat walls, biofilms could provide dynamic functionality by regulating humidity and gas exchange. Their incorporation could enable engineered amyloid or polysaccharide matrices to act as living sensors, reporting structural fatigue, detecting chemical leaks or modulating material behaviour in response to environmental cues. In effect, these living layers would provide adaptable, self-maintaining protection.

In terms of life support and power systems, biofilm engineering has the potential to revolutionise both. The ability to program division of labour and metabolic complementarity into biofilms make them ideal candidates for modular bioprocessing units which can deliver both power and waste bioremediation concomitantly, as is evidenced in current microbial fuel cell development (Gao et al., 2024; Mahto and Das, 2025; Ye et al., 2025). With bioregenerative life support systems (BLSS), an integrated approach is required, as described in reviews by De Micco et al. (2023) and Koehle et al. (2023). Biofilm communities could be engineered for optimal CO2 processing and O2 production, water filtration and detoxification of spacecraft atmospheres, somewhat akin to lung, kidney and liver functionality. Designing spatially structured biofilms produced via lithography or seeded 3D architectures would allow fine-tuned control over metabolic and waste processing zones, enabling efficient cross-feeding and syntrophic interactions essential for stable long-term operation.

Biofilm engineering will also be essential for the integration of settlements and the in-situ resource utilisation beyond Earth. There is much study on the biomining and bioleaching of materials from regolith (Vezzola et al., 2023; Flores et al., 2025; Figuera et al., 2025) and for sustainable mining and bioremediation in both Terran and non-Terran environments (Cockell et al., 2021; Mishra et al., 2022; Majhi et al., 2024; Savadiya et al., 2024; Cozma et al., 2024; Mathivanan et al., 2025; Santomartino et al., 2026). Although Earth-based research shows the ability of interkingdom microbial biofilms to bioremediate soil contaminants (Amir et al., 2026), their behaviour in regolith is yet to be addressed. These biomining and bioremediation capacities position biofilms as true pioneer systems. Engineered biofilms offer resource generation for colony establishment and provide capacity for initiation of ecological succession in the wider extraterrestrial environment by stabilising regolith and creating localised microenvironments for other species. Furthermore, as integral components of organism and environmental wellbeing, engineering, and augmentation of health-associated biofilms in both crew and plants will be essential to counteract spaceflight-induced health conditions (Baxter et al., 2026).

5 Conclusion

Viewing biofilms as rudimentary tissue provides a coherent foundation for engineering them as living materials. Although current technologies allow manipulation of cells, matrix components and spatial organisation, these approaches remain fragmented. Achieving predictable, whole-biofilm engineering will require deeper mechanistic insight into community interactions, environmental feedback loops and gravity-dependent biology, alongside new modelling frameworks and tools that operate at the community scale.

The implications for space exploration are substantial as engineered biofilms offer capabilities unmatched by abiotic systems. Their capacity for self-assembly, self-repair, metabolic breadth, and environmental adaptability position biofilms as astro-pioneer systems, living technologies capable of initiating, stabilising, and transforming extraterrestrial environments. From radiation-attenuating composites and regolith-binding materials to bioregenerative life support, microbial power systems and health countermeasures, biofilms could underpin many of the biological functions and crew support required for long-duration missions and off-world settlement. Advancing whole-biofilm engineering will therefore be central to realising their full potential as adaptive systems for life beyond Earth.

Generative AI statement

The author(s) declared that generative AI was used in the creation of this manuscript. Microsoft Copilot (Microsoft Corporation) was used for sentence optimisation and critique of the manuscript. Prompts used for this purpose are included in Supplementary Material.

Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.

Publisher’s note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

Supplementary material

The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fspas.2026.1843790/full#supplementary-material

Supplementary material Supplementary material at the source

Footnotes

Edited by: Daniela Billi, University of Rome Tor Vergata, Italy

Reviewed by: Clay Fuqua, Indiana University Bloomington, United States

Pamela Flores, Space Lab Technologies, United States

Tamanna Jafar, University of Tennessee, United States

Statements

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Funding

The author(s) declared that financial support was received for this work and/or its publication. K.J.B was funded by the Biotechnology and Biological Sciences Research Council (BBSRC) Grant number BB/Z516120/1.

Data availability

This is a Mini Review article with no new data created or modified, and therefore all data underlying this study is cited in the references.

Author contributions

KJB: Conceptualization, Investigation, Writing – original draft, Writing – review and editing.

References

  1. Abbott R. D., Koptiuch C., Iatridis J. C., Howe A. K., Badger G. J., Langevin H. M. (2013). Stress and matrix‐responsive cytoskeletal remodeling in fibroblasts. J. Cellular Physiology 228 (1), 50 – 57. doi:10.1002/jcp.24102
  2. Abdel-Nour M., Su H., Duncan C., Li S., Raju D., Shamoun F. (2019). Polymorphisms of a collagen-like adhesin contributes to Legionella pneumophila adhesion, biofilm formation capacity and clinical prevalence. Front. Microbiol. 10, 604. doi:10.3389/fmicb.2019.00604
  3. Aguilar C., Eichwald C., Eberl L. (2015). “ Multicellularity in bacteria: from division of labor to biofilm formation,” in Evolutionary Transitions to Multicellular Life: Principles and Mechanisms (Dordrecht: Springer), 79 – 95. doi:10.1007/978-94-017-9642-2_4
  4. Aliyazdi S., Frisch S., Hidalgo A., Frank N., Krug D., Müller R. (2023). 3D bioprinting of E. coli MG1655 biofilms on human lung epithelial cells for building complex in vitro infection models. Biofabrication 15 (3), 035019. doi:10.1088/1758-5090/acd95e
  5. Amir H., Guentas L., Crossay T., Bourles A., Burtet-Sarramegna V. (2026). Synergism of arbuscular mycorrhizal fungi and bacteria in bioremediation and restoration of metal-stressed environments. Front. Plant Sci. 17, 1753034. doi:10.3389/fpls.2026.1753034
  6. Azevedo A., Teixeira-Santos R., Gomes L. C., Duarte S. O., Monteiro G. A., Mergulhão F. J. (2025). Engineering Escherichia coli biofilms for curcumin production. Molecules 30 (9), 2031. doi:10.3390/molecules30092031
  7. Bakhshandeh B., Ranjbar N., Abbasi A., Amiri E., Abedi A., Mehrabi M. R. (2023). Recent progress in the manipulation of biochemical and biophysical cues for engineering functional tissues. Bioeng. & Translational Medicine 8 (2), e10383. doi:10.1002/btm2.10383
  8. Baxter K. J., Sas E., Clark K. B., Walsh M., Pradeep N., Batool A. (2026). Biofilms: from the cradle of life to life support. Npj Biofilms Microbiomes 12 (1), 11. doi:10.1038/s41522-025-00875-8
  9. Bjarnsholt T., Whiteley M., Rumbaugh K. P., Stewart P. S., Jensen P. Ø., Frimodt-Møller N. (2022). The importance of understanding the infectious microenvironment. Lancet Infect. Dis. 22 (3), e88 – e92. doi:10.1016/S1473-3099(21)00122-5
  10. Bratkic A., Jazbec A., Toplak N., Koren S., Lojen S., Tinta T. (2024). The colonization of an irradiated environment: the case of microbial biofilm in a nuclear reactor. Int. Journal Radiation Biology 100 (1), 108 – 121. doi:10.1080/09553002.2023.2258206
  11. Cao X., Wang X., Chen R., Chen L., Liu Y., Wang M. (2025). Improving Bacillus subtilis as biological chassis performance by the CRISPR genetic toolkit. ACS Synth. Biol. 14 (3), 677 – 688. doi:10.1021/acssynbio.4c00844
  12. Chai M., Deng C., Chen Q., Lu W., Liu Y., Li J. (2021). Synthetic biology toolkits and metabolic engineering applied in Corynebacterium glutamicum for biomanufacturing. ACS Synth. Biol. 10 (12), 3237 – 3250. doi:10.1021/acssynbio.1c00355
  13. Chen Z., Srivastava P., Zarazúa-Osorio B., Marathe A., Fujita M., Igoshin O. A. (2022). Bacillus subtilis histidine kinase KinC activates biofilm formation by controlling heterogeneity of single-cell responses. MBio 13 (1), e01694. doi:10.1128/mbio.01694-21
  14. Chen G. Q., Zhang X., Liu X., Huang W., Xie Z., Han J. (2022). H alomonas spp., as chassis for low-cost production of chemicals. Appl. Microbiol. Biotechnol. 106 (21), 6977 – 6992. doi:10.1007/s00253-022-12215-3
  15. Cockell C. S., Santomartino R., Finster K., Waajen A. C., Nicholson N., Loudon C. M. (2021). Microbially-enhanced vanadium mining and bioremediation under micro-and Mars gravity on the International Space station. Front. Microbiology 12, 641387. doi:10.3389/fmicb.2021.641387
  16. Cooke M. B., Welch K. T., Ramirez L. D., Wen A. X., Marciano D. C., Herman C. (2025). GoldenBraid2. 0 E. coli: a comprehensive and characterized toolkit for enterics. Synth. Biol. 10 (1), ysaf015. doi:10.1093/synbio/ysaf015
  17. Cozma P., Bețianu C., Hlihor R. M., Simion I. M., Gavrilescu M. (2024). Bio-recovery of metals through biomining within circularity-based solutions. Processes 12 (9), 1793. doi:10.3390/pr12091793
  18. De Belly H., Paluch E. K., Chalut K. J. (2022). Interplay between mechanics and signalling in regulating cell fate. Nat. Rev. Mol. Cell Biol. 23 (7), 465 – 480. doi:10.1038/s41580-022-00472-z
  19. De Micco V., Amitrano C., Mastroleo F., Aronne G., Battistelli A., Carnero-Diaz E. (2023). Plant and microbial science and technology as cornerstones to Bioregenerative Life Support Systems in space. Npj Microgravity 9 (1), 69. doi:10.1038/s41526-023-00317-9
  20. Della Valle S., Tu W., Huang W. E. (2024). Construction of microbial platform chassis for CO2 utilisation. Curr. Opin. Syst. Biol. 37, 100489. doi:10.1016/j.coisb.2023.100489
  21. Denton O., Wan Y., Beattie L., Jack T., McGoldrick P., McAllister H. (2024). Understanding the role of biofilms in acute recurrent tonsillitis through 3D bioprinting of a novel gelatin-PEGDA hydrogel. Bioengineering 11 (3), 202. doi:10.3390/bioengineering11030202
  22. Deveau A., Bonito G., Uehling J., Paoletti M., Becker M., Bindschedler S. (2018). Bacterial–fungal interactions: ecology, mechanisms and challenges. FEMS Microbiology Reviews 42 (3), 335 – 352. doi:10.1093/femsre/fuy008
  23. DiCandia M. A., Edwards A. N., Alcaraz Y. B., Monteiro M. P., Lee C. D., Vargas Cuebas G. (2024). A conserved switch controls virulence, sporulation, and motility in C. difficile. PLoS Pathog. 20 (5), e1012224. doi:10.1371/journal.ppat.1012224
  24. Dong H., Zhang W., Wang Y., Liu D., Wang P. (2020). Biofilm polysaccharide display platform: a natural, renewable, and biocompatible material for improved lipase performance. J. Agricultural Food Chemistry 68 (5), 1373 – 1381. doi:10.1021/acs.jafc.9b07209
  25. Doupé D. P., Perrimon N. (2016). Toward a systems understanding of signaling pathway function. Curr. Top. Dev. Biol. 117, 221 – 236. doi:10.1016/bs.ctdb.2015.11.005
  26. Duraj-Thatte A. M., Manjula-Basavanna A., Courchesne N. M. D., Cannici G. I., Sánchez-Ferrer A., Frank B. P. (2021). Water-processable, biodegradable and coatable aquaplastic from engineered biofilms. Nat. Chemical Biology 17 (6), 732 – 738. doi:10.1038/s41589-021-00773-y
  27. Durmusoglu D., Al’Abri I., Li Z., Islam Williams T., Collins L. B., Martínez J. L. (2023). Improving therapeutic protein secretion in the probiotic yeast Saccharomyces boulardii using a multifactorial engineering approach. Microb. Cell Factories 22 (1), 109. doi:10.1186/s12934-023-02117-y
  28. Ertelt M. J., Eicheler C., Machner A., Lieleg O. (2022). Durability of biofilm-enriched hybrid mortar towards chemical and physical challenges. Mater. Lett. 326, 132937. doi:10.1016/j.matlet.2022.132937
  29. Espina J. A., Cordeiro M. H., Milivojevic M., Pajić-Lijaković I., Barriga E. H. (2023). Response of cells and tissues to shear stress. J. Cell Sci. 136 (18), jcs260985. doi:10.1242/jcs.260985
  30. Espinoza Miranda S. S., Abbaszade G., Hess W. R., Drescher K., Saliba A. E., Zaburdaev V. (2025). Resolving spatiotemporal dynamics in bacterial multicellular populations: approaches and challenges. Microbiol. Mol. Biol. Rev. 89 (1), e00138. doi:10.1128/mmbr.00138-24
  31. Faber A., Politan R. J., Stukenberg D., Morris K. M., Kim R., Jeon E. (2025). Expanding genetic engineering capabilities in Vibrio natriegens with the Vnat Collection. Nucleic Acids Res. 53 (13), gkaf580. doi:10.1093/nar/gkaf580
  32. Fede C., Pirri C., Fan C., Petrelli L., Guidolin D., De Caro R. (2021). A closer look at the cellular and molecular components of the deep/muscular fasciae. Int. Journal Molecular Sciences 22 (3), 1411. doi:10.3390/ijms22031411
  33. Felz S., Neu T. R., van Loosdrecht M. C., Lin Y. (2020). Aerobic granular sludge contains hyaluronic acid-like and sulfated glycosaminoglycans-like polymers. Water Research 169, 115291. doi:10.1016/j.watres.2019.115291
  34. Figueira J., Koch S., Müller D. W., Slawik S., Cowley A., Moeller R. (2025). Biomining of lunar regolith simulant EAC-1 A with the fungus Penicillium simplicissimum. Fungal Biol. Biotechnol. 12 (1), 8. doi:10.1186/s40694-025-00201-z
  35. Flemming H. C., van Hullebusch E. D., Neu T. R., Nielsen P. H., Seviour T., Stoodley P. (2023). The biofilm matrix: multitasking in a shared space. Nat. Rev. Microbiol. 21 (2), 70 – 86. doi:10.1038/s41579-022-00791-0
  36. Flores P., Escobar A., Garcia N. D., Chamberlain C. (2025). EcoMine™-Bioregenerative Mineral Mining from Lunar Regolith. doi:10.32865/2346/102876
  37. Gao X., Sun T., Pei G., Chen L., Zhang W. (2016). Cyanobacterial chassis engineering for enhancing production of biofuels and chemicals. Appl. Microbiology Biotechnology 100 (8), 3401 – 3413. doi:10.1007/s00253-016-7374-2
  38. Gao S. C., Fan X. X., Zhang Z., Li R. T., Zhang Y., Gao T. P. (2024). A dual-function mixed-culture biofilm for sulfadiazine removal and electricity production using bio-electrochemical system. Biosens. Bioelectron. 263, 116552. doi:10.1016/j.bios.2024.116552
  39. González-Rojo S., Paniagua-García A. I., Díez-Antolínez R. (2024). Advances in microbial biotechnology for sustainable alternatives to petroleum-based plastics: a comprehensive review of polyhydroxyalkanoate production. Microorganisms 12 (8), 1668. doi:10.3390/microorganisms12081668
  40. Gudipaty S. A., Lindblom J., Loftus P. D., Redd M. J., Edes K., Davey C. F. (2017). Mechanical stretch triggers rapid epithelial cell division through Piezo1. Nature 543 (7643), 118 – 121. doi:10.1038/nature21407
  41. Heino J. (2014). Cellular signaling by collagen-binding integrins. I Domain Integrins 819, 143 – 155. doi:10.1007/978-94-017-9153-3_10
  42. Huang J., Liu S., Zhang C., Wang X., Pu J., Ba F. (2019). Programmable and printable Bacillus subtilis biofilms as engineered living materials. Nat. Chemical Biology 15 (1), 34 – 41. doi:10.1038/s41589-018-0169-2
  43. Jin X., Riedel-Kruse I. H. (2018). Biofilm Lithography enables high-resolution cell patterning via optogenetic adhesin expression. Proc. Natl. Acad. Sci. 115 (14), 3698 – 3703. doi:10.1073/pnas.1720676115
  44. Jin X., Riedel-Kruse I. H. (2024). Optogenetic patterning generates multi-strain biofilms with spatially distributed antibiotic resistance. Nat. Commun. 15 (1), 9443. doi:10.1038/s41467-024-53546-1
  45. Jo J., Price-Whelan A., Dietrich L. E. (2022). Gradients and consequences of heterogeneity in biofilms. Nat. Rev. Microbiol. 20 (10), 593 – 607. doi:10.1038/s41579-022-00692-2
  46. Johnston T. G., Yuan S. F., Wagner J. M., Yi X., Saha A., Smith P. (2020). Compartmentalized microbes and co-cultures in hydrogels for on-demand bioproduction and preservation. Nat. Communications 11 (1), 563. doi:10.1038/s41467-020-14371-4
  47. Karim M. A., LaJeunesse D. (2026). Acute High Shear Stress Enhances Fungal cell/substrate Adhesion 14 (4), e01952. doi:10.1128/spectrum.01952-25
  48. Kim B., Madukoma C. S., Shrout J. D., Nerenberg R. (2023). Effect of EPS production on the performance of membrane-based biofilm reactors. Water Res. 240, 120101. doi:10.1016/j.watres.2023.120101
  49. Knott S., Curry D., Zhao N., Metgud P., Dastgheyb S. S., Purtill C. (2021). Staphylococcus aureus floating biofilm formation and phenotype in synovial fluid depends on albumin, fibrinogen, and hyaluronic acid. Front. Microbiol. 12, 655873. doi:10.3389/fmicb.2021.655873
  50. Kochina T. A., Kondratenko Y. A., Shilova O. A., Vlasov D. Y. (2022). Biocorrosion, biofouling, and advanced methods of controlling them. Prot. Metals Phys. Chem. Surfaces 58 (1), 129 – 150. doi:10.1134/S2070205122010129
  51. Koehle A. P., Brumwell S. L., Seto E. P., Lynch A. M., Urbaniak C. (2023). Microbial applications for sustainable space exploration beyond low Earth orbit. Npj Microgravity 9 (1), 47. doi:10.1038/s41526-023-00285-0
  52. Kumari S., Siingh S., Priyadarshanee M., Das S. (2025). Functional amyloid fibrils in the marine bacterium Pseudomonas aeruginosa PFL-P1 interact with lead and upregulate the fapC gene for heavy metal sequestration. J. Environ. Chem. Eng. 13 (3), 116235. doi:10.1016/j.jece.2025.116235
  53. Kurakado S., Arai R., Sugita T. (2018). Association of the hypha‐related protein Pra1 and zinc transporter Zrt1 with biofilm formation by the pathogenic yeast Candida albicans. Microbiol. Immunol. 62 (6), 405 – 410. doi:10.1111/1348-0421.12596
  54. Li F., Li X., Ye L., Liu X., Zhu J., Yang S. (2022). A genetically engineered composite biofilm for microbial induced calcium carbonate precipitation by synergic effect of urease, protein adhesive and xanthan gum. J. Environ. Chem. Eng. 10 (5), 108431. doi:10.1016/j.jece.2022.108431
  55. Li Z., Ren Y., Li Z., Zhang J., Fan Y., Jiang G. (2024). Engineered living biofilm with enhanced metal binding ability for corrosion protection in seawater. Adv. Funct. Mater. 34 (13), 2313120. doi:10.1002/adfm.202313120
  56. Liang Y., Motawaa M., Bu X., Wei J., Shao Y., Li Y. (2025). Construction of primary chassis cells with efficient protein expression in Thermus thermophilus. Microb. Cell Factories 24 (1), 163. doi:10.1186/s12934-025-02785-y
  57. Lichtenberg M., Kragh K. N., Fritz B., Kirkegaard J. B., Tolker-Nielsen T., Bjarnsholt T. (2022). Cyclic-di-GMP signaling controls metabolic activity in Pseudomonas aeruginosa. Cell Reports 41 (3), 111515. doi:10.1016/j.celrep.2022.111515
  58. Lipke P. N. (2025). Not gently down the stream: flow induces amyloid bonding in environmental and pathological fungal biofilms. Mbio 16 (6), e00203 – e00225. doi:10.1128/mbio.00203-25
  59. Liu X., Han X., Peng Y., Tan C., Wang J., Xue H. (2022). Rapid production of l‐DOPA by Vibrio natriegens, an emerging next‐generation whole‐cell catalysis chassis. Microb. Biotechnol. 15 (5), 1610 – 1621. doi:10.1111/1751-7915.14001
  60. Long Y., Han X., Meng X., Xu P., Tao F. (2024). A robust yeast chassis: comprehensive characterization of a fast-growing Saccharomyces cerevisiae. Mbio 15 (2), e0319. doi:10.1128/mbio.03196-23
  61. Mahto K. U., Das S. (2025). Electroactive biofilm communities in microbial fuel cells for the synergistic treatment of wastewater and bioelectricity generation. Crit. Rev. Biotechnol. 45 (2), 434 – 453. doi:10.1080/07388551.2024.2372070
  62. Maillard J. Y., Centeleghe I. (2023). How biofilm changes our understanding of cleaning and disinfection. Antimicrob. Resist. Infect. Control 12 (1), 95. doi:10.1186/s13756-023-01290-4
  63. Majhi B., Semwal P., Mishra S. K., Chauhan P. S. (2024). Strategies for microbes-mediated arsenic bioremediation: impact of quorum sensing in the rhizosphere. Sci. Total Environ. 956, 177321. doi:10.1016/j.scitotenv.2024.177321
  64. Mao Y., Wickström S. A. (2024). Mechanical state transitions in the regulation of tissue form and function. Nat. Rev. Mol. Cell Biol. 25 (8), 654 – 670. doi:10.1038/s41580-024-00719-x
  65. Martinez-Garcia E., de Lorenzo V. (2024). Pseudomonas putida as a synthetic biology chassis and a metabolic engineering platform. Curr. Opin. Biotechnol. 85, 103025. doi:10.1016/j.copbio.2023.103025
  66. Mathivanan K., Zhang R., Chandirika J. U., Mathimani T., Wang C., Duan J. (2025). Bacterial biofilm-based bioleaching: sustainable mitigation and potential management of e-waste pollution. Waste Manag. 193, 221 – 236. doi:10.1016/j.wasman.2024.12.010
  67. Matilla-Cuenca L., Taglialegna A., Gil C., Toledo-Arana A., Lasa I., Valle J. (2022). Bacterial biofilm functionalization through Bap amyloid engineering. NPJ Biofilms Microbiomes 8 (1), 62. doi:10.1038/s41522-022-00324-w
  68. Matsumoto T., Tanaka T., Kondo A. (2017). Engineering metabolic pathways in Escherichia coli for constructing a “microbial chassis” for biochemical production. Bioresour. Technology 245, 1362 – 1368. doi:10.1016/j.biortech.2017.05.008
  69. Meng Q., Xia Y. (2011). c-Jun, at the crossroad of the signaling network. Protein & Cell 2 (11), 889 – 898. doi:10.1007/s13238-011-1113-3
  70. Mishra S., Huang Y., Li J., Wu X., Zhou Z., Lei Q. (2022). Biofilm-mediated bioremediation is a powerful tool for the removal of environmental pollutants. Chemosphere 294, 133609. doi:10.1016/j.chemosphere.2022.133609
  71. Nguyen P. Q., Botyanszki Z., Tay P. K. R., Joshi N. S. (2014). Programmable biofilm-based materials from engineered curli nanofibres. Nat. Communications 5 (1), 4945. doi:10.1038/ncomms5945
  72. Nikolaev Y. A., Plakunov V. K. (2007). Biofilm—“City of microbes” or an analogue of multicellular organisms? Microbiology 76 (2), 125 – 138. doi:10.1134/S0026261707020014
  73. Pan R., Yuan Y., Xu A., Jiang W., Zhang W., Barriuso J. (2025). Biofilm engineering to improve succinic acid production in Escherichia coli by enhancing extracellular polysaccharide synthesis. Bioresour. Technol. 431, 132627. doi:10.1016/j.biortech.2025.132627
  74. Penesyan A., Paulsen I. T., Kjelleberg S., Gillings M. R. (2021). Three faces of biofilms: a microbial lifestyle, a nascent multicellular organism, and an incubator for diversity. Npj Biofilms Microbiomes 7 (1), 80. doi:10.1038/s41522-021-00251-2
  75. Percival S. L., Suleman L., Vuotto C., Donelli G. (2015). Healthcare-associated infections, medical devices and biofilms: risk, tolerance and control. J. Medical Microbiology 64 (4), 323 – 334. doi:10.1099/jmm.0.000032
  76. Pisithkul T., Schroeder J. W., Trujillo E. A., Yeesin P., Stevenson D. M., Chaiamarit T. (2019). Metabolic remodeling during biofilm development of Bacillus subtilis. MBio 10 (3), 10 – 1128. doi:10.1128/mBio.00623-19
  77. Poohphajai F., Gubenšek A., Černoša A., Butina Ogorelec K., Rautkari L., Sandak J. (2024). Bioinspired living coating System for wood protection: exploring fungal species on wood surfaces coated with biofinish during its service life. Coatings 14 (4), 430. doi:10.3390/coatings14040430
  78. Prasad B., Richter P., Vadakedath N., Haag F. W., Strauch S. M., Mancinelli R. (2021). How the space environment influences organisms: an astrobiological perspective and review. Int. J. Astrobiol. 20 (2), 159 – 177. doi:10.1017/S1473550421000057
  79. Praveschotinunt P., Duraj-Thatte A. M., Gelfat I., Bahl F., Chou D. B., Joshi N. S. (2019). Engineered E. coli Nissle 1917 for the delivery of matrix-tethered therapeutic domains to the gut. Nat. Communications 10 (1), 5580. doi:10.1038/s41467-019-13336-6
  80. Purcell E. B., McKee R. W., Courson D. S., Garrett E. M., McBride S. M., Cheney R. E. (2017). A nutrient-regulated cyclic diguanylate phosphodiesterase controls Clostridium difficile biofilm and toxin production during stationary phase. Infect. Immunity 85 (9), 10 – 1128. doi:10.1128/IAI.00347-17
  81. Quan T., Qin Z., He T., Fisher G. J. (2025). Integrin α11β1 as a key collagen receptor in human skin dermis: insight into fibroblast function and skin dermal aging. J. Investigative Dermatology 145 (10), 2449 – 2463. doi:10.1016/j.jid.2025.03.006
  82. Rados T., Leland O. S., Escudeiro P., Mallon J., Andre K., Caspy I. (2025). Tissue-like multicellular development triggered by mechanical compression in archaea. Science 388 (6742), 109 – 115. doi:10.1126/science.adu0047
  83. Ragon M., Restoux G., Moreira D., Møller A. P., López-García P. (2011). Sunlight-exposed biofilm microbial communities are naturally resistant to Chernobyl ionizing-radiation levels. PloS One 6 (7), e21764. doi:10.1371/journal.pone.0021764
  84. Rahayu D. S., Ambarsari L., Nurcholis W., Shalsabilla S. E., Fadilah A., Sanjaya M. F. (2025). Microbial diversity of biofilm in spent nuclear fuel storage pond. J. Appl. Biol. & Biotechnol. 13 (1), 143 – 149. doi:10.7324/JABB.2024.194708
  85. Răut I., Călin M., Vuluga Z., Oancea F., Paceagiu J., Radu N. (2021). Fungal based biopolymer composites for construction materials. Materials 14 (11), 2906. doi:10.3390/ma14112906
  86. Ren P., Zhao W., Zhou C., Chen T., Sun W., Chen Y. (2025). Green and sustainable FAME production: enhanced continuous catalysis based on biofilm surface-displayed enzymes. Algal Res. 88, 103984. doi:10.1016/j.algal.2025.103984
  87. Rossi C. S., Eckartt K., Scarchilli E., Angeli S., Price-Whelan A., Di Matteo A. (2023). Molecular insights into RmcA-mediated c-di-GMP consumption: linking redox potential to biofilm morphogenesis in Pseudomonas aeruginosa. Microbiol. Res. 277, 127498. doi:10.1016/j.micres.2023.127498
  88. Sadiq F. A., Hansen M. F., Burmølle M., Heyndrickx M., Flint S., Lu W. (2022). Trans-kingdom interactions in mixed biofilm communities. FEMS Microbiology Reviews 46 (5), fuac024. doi:10.1093/femsre/fuac024
  89. Santomartino R., Rodriguez Blanco G., Gudgeon A., Hafner J. H., Stirpe A., Waterfall M. (2026). Microbial biomining from asteroidal material onboard the international space station. Microgravity. 23. doi:10.1038/s41526-026-00567-3
  90. Savadiya B., Pandey G., Misra S. K. (2024). Remediation of pharmacophoric laboratory waste by using biodegradable carbon nanoparticles of bacterial biofilm origin. Environ. Res. 252, 118969. doi:10.1016/j.envres.2024.118969
  91. Schmieden D. T., Basalo Vázquez S. J., Sanguesa H., Van Der Does M., Idema T., Meyer A. S. (2018). Printing of patterned, engineered E. coli biofilms with a low-cost 3D printer. ACS Synthetic Biology 7 (5), 1328 – 1337. doi:10.1021/acssynbio.7b00424
  92. Senatore G., Mastroleo F., Leys N., Mauriello G. (2018). Effect of microgravity & space radiation on microbes. Future Microbiology 13 (7), 831 – 847. doi:10.2217/fmb-2017-0251
  93. Serra D. O., Hengge R. (2014). Stress responses go three dimensional–the spatial order of physiological differentiation in bacterial macrocolony biofilms. Environ. Microbiology 16 (6), 1455 – 1471. doi:10.1111/1462-2920.12483
  94. Shrestha P., Kim B., Han S. R., Lee H., Oh T. J. (2025). Unraveling the genetic mechanisms of UV radiation resistance in bacillus through biofilm formation, sporulation, and carotenoid production. Genomics 117 (4), 111066. doi:10.1016/j.ygeno.2025.111066
  95. Simsek M. F., Özbudak E. M. (2022). Patterning principles of morphogen gradients. Open Biol. 12 (10), 220224. doi:10.1098/rsob.220224
  96. Soloviev D. A., Fonzi W. A., Sentandreu R., Pluskota E., Forsyth C. B., Yadav S. (2007). Identification of pH-regulated antigen 1 released from Candida albicans as the major ligand for leukocyte integrin αMβ2. J. Immunology 178 (4), 2038 – 2046. doi:10.4049/jimmunol.178.4.2038
  97. Sun D., Liu X., Zhang Y., Shi R., Ru Y., Zhou X. (2025). Local c-di-GMP signaling, triggered by cross-regulation of cAMP-CRP and c-di-GMP, controls biofilm formation under nutrient limitation. Proc. Natl. Acad. Sci. 122 (35), e2516964122. doi:10.1073/pnas.2516964122
  98. Tay P. K. R., Nguyen P. Q., Joshi N. S. (2017). A synthetic circuit for mercury bioremediation using self-assembling functional amyloids. ACS Synthetic Biology 6 (10), 1841 – 1850. doi:10.1021/acssynbio.7b00137
  99. Váchová L., Plocek V., Maršíková J., Rešetárová S., Hatáková L., Palková Z. (2024). Differential stability of Gcn4p controls its cell-specific activity in differentiated yeast colonies. MBio 15 (5), e00689. doi:10.1128/mbio.00689-24
  100. Van Gestel J., Vlamakis H., Kolter R. (2015). From cell differentiation to cell collectives: bacillus subtilis uses division of labor to migrate. PLoS Biology 13 (4), e1002141. doi:10.1371/journal.pbio.1002141
  101. Vezzola M., Tosi S., Doria E., Bonazzi M., Alvaro M., Sanfilippo A. (2023). Interaction between a Martian regolith simulant and fungal organic acids in the biomining perspective. J. Fungi 9 (10), 976. doi:10.3390/jof9100976
  102. Vijayakumar V. E., Venkataraman K. (2024). A systematic review of the potential of Pichia pastoris (Komagataella phaffii) as an alternative host for biologics production. Mol. Biotechnol. 66 (7), 1621 – 1639. doi:10.1007/s12033-023-00803-1
  103. Wang X., Zhang J., Li K., An B., Wang Y., Zhong C. (2022). Photocatalyst-mineralized biofilms as living bio-abiotic interfaces for single enzyme to whole-cell photocatalytic applications. Sci. Adv. 8 (18), eabm7665. doi:10.1126/sciadv.abm7665
  104. Wei Y., Zhou X., Li Z., Liu Q., Ding H., Zhou Y. (2025). Genetically programmed single‐component protein hydrogel for Spinal Cord Injury Repair. Adv. Sci. 12 (10), 2405054. doi:10.1002/advs.202405054
  105. Wong L. L., Mugunthan S., Kundukad B., Ho J. C. S., Rice S. A., Hinks J. (2023). Microbial biofilms are shaped by the constant dialogue between biological and physical forces in the extracellular matrix. Environ. Microbiol, 199 – 208. doi:10.1111/1462-2920.16306
  106. Xue Y., Kong C., Shen W., Bai C., Ren Y., Zhou X. (2017). Methylotrophic yeast Pichia pastoris as a chassis organism for polyketide synthesis via the full citrinin biosynthetic pathway. J. Biotechnology 242, 64 – 72. doi:10.1016/j.jbiotec.2016.11.031
  107. Xue C., Chu Q., Shi Q., Zeng Y., Lu J., Li L. (2025). Wnt signaling pathways in biology and disease: mechanisms and therapeutic advances. Signal Transduct. Target. Ther. 10 (1), 106. doi:10.1038/s41392-025-02142-w
  108. Yadav M. K., Chae S. W., Park K., Song J. J. (2013). Hyaluronic acid derived from other streptococci supports Streptococcus pneumoniae in vitro biofilm formation. BioMed Research International 2013 (1), 690217. doi:10.1155/2013/690217
  109. Yaman Y. I., Demir E., Vetter R., Kocabas A. (2019). Emergence of active nematics in chaining bacterial biofilms. Nat. Communications 10 (1), 2285. doi:10.1038/s41467-019-10311-z
  110. Yang M., Hutchinson N., Ye N., Timek H., Jennings M., Yin J. (2024). Engineered Bacillus subtilis as oral probiotics to enhance clearance of blood lactate. ACS Synthetic Biology 14 (1), 101 – 112. doi:10.1021/acssynbio.4c00399
  111. Ye J., Gu W., Hu J., Chen L., Yang C., Gao J. (2025). Toward next-generation semiartificial photosynthesis: multidisciplinary engineering of biohybrid systems. Chem. Rev. 125 (24), 12198 – 12252. doi:10.1021/acs.chemrev.5c00658
  112. Zhang J., Wu H., Wang D., Wang L., Cui Y., Zhang C. (2022). Intracellular glycosyl hydrolase PslG shapes bacterial cell fate, signaling, and the biofilm development of Pseudomonas aeruginosa. Elife 11, e72778. doi:10.7554/eLife.72778
  113. Zhang L., Lin Y., Yi X., Huang W., Hu Q., Zhang Z. (2023). Engineering low-salt growth Halomonas bluephagenesis for cost-effective bioproduction combined with adaptive evolution. Metab. Eng. 79, 146 – 158. doi:10.1016/j.ymben.2023.08.001
  114. Zhao X., Wang Y., Shang Q., Li Y., Hao H., Zhang Y. (2015). Collagen-like proteins (ClpA, ClpB, ClpC, and ClpD) are required for biofilm formation and adhesion to plant roots by Bacillus amyloliquefaciens FZB42. PloS One 10 (2), e0117414. doi:10.1371/journal.pone.0117414
  115. Zhu B., Ye Q., Seo Y., Wei N. (2022). Enzymatic degradation of polyethylene terephthalate plastics by bacterial curli display PETase. Environ. Sci. & Technol. Lett. 9 (7), 650 – 657. doi:10.1021/acs.estlett.2c00332

Citation

Baxter, K. J. (2026). Building biofilm: engineering for life off Earth. Frontiers in Astronomy and Space Sciences, Volume 13. https://doi.org/
Received
Accepted
Published
Volume
13 - 2026

Copyright

Copyright © 2026 Baxter.

License: https://creativecommons.org/licenses/by/4.0/

*Correspondence: Katherine J. Baxter

Article metrics

Views
0
Downloads
0
Citations
0
Shares
0

Related articles

Original ResearchProperties of the circumstellar matter around the pair-instability supernova candidate SN 2018ibb revealed by its light curveTakashi J. Moriya, Steve Schulze, Alexander Heger, Sergei I. Blinnikov, Marat Sh. PotashovOriginal ResearchCrowdsourced aurora observations during geomagnetic superstorms: the Aurorasaurus open science databaseElizabeth MacDonald, Feras Natsheh, Laura Brandt Edson, Matthew HeavnerOriginal ResearchNewtonian gravity and dark-matter phenomenology from an emergent scalar field sourced by ordinary matterGustav Stålhammar