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Review

Algicidal Bacteria: A Sustainable Proposal to Control Microalgae in the Conservation and Restoration of Stone Cultural Heritage

by
Isabel Calvo-Bayo
1,
Fernando Bolívar-Galiano
1 and
Julio Romero-Noguera
2,*
1
Painting Department, Faculty of Fine Arts, University of Granada, Avda. Andalucía s/n, 18071 Granada, Spain
2
Painting Department, Faculty of Fine Arts, University of Seville, Laraña 3, 41003 Seville, Spain
*
Author to whom correspondence should be addressed.
Sustainability 2025, 17(23), 10610; https://doi.org/10.3390/su172310610
Submission received: 12 September 2025 / Revised: 13 November 2025 / Accepted: 21 November 2025 / Published: 26 November 2025
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Abstract

The growth of microalgae poses a significant threat to the preservation of stone heritage, particularly in ornamental fountains and water-related architecture. Traditional chemical cleaning methods, such as quaternary ammonium compounds and chlorine-based solutions, are often ineffective and can be harmful to both the environment and cultural properties. In response, biocleaning, which involves the use of live microorganisms and is part of biorestoration, is gaining prominence in cultural heritage conservation, offering a sustainable alternative to conventional methods. The use of microorganisms antagonistic to microalgae growth has been extensively studied in environmental biotechnology to eliminate harmful algae, though its application in heritage conservation remains limited. This review summarizes current knowledge on bacteria capable of inhibiting microalgae growth, discussing their mechanisms, effectiveness, and potential applications, alongside the environmental and economic benefits and challenges of these methods. By collating and critically assessing available information, this paper aims to serve as a comprehensive resource for conservators, restorers, and researchers interested in innovative and sustainable approaches to combat biodeterioration in stone heritage, thereby fostering the development of effective and environmentally sustainable treatments for such culturally significant properties.

1. Introduction

Algal overgrowth presents a significant challenge in preserving cultural heritage sites, particularly those whose main constituent material is natural stone. Microalgae can thrive in nutrient-poor environments such as stone surfaces and proliferate rapidly in areas with high humidity or continuous water presence. Their development is especially problematic in fountains and ornamental pools, where water exposure creates ideal growth conditions (Figure 1) [1,2].
Microalgae, along with fungi and bacteria, are primary agents of biodeterioration, causing a wide range of esthetic, physical, and chemical damage to stone cultural assets (Figure 2). This includes discoloration, weakening, and structural degradation, all extensively documented worldwide, highlighting the need for effective and sustainable control measures. Addressing this issue is crucial for the long-term preservation of significant cultural heritage [3,4,5,6].
Photosynthetic microorganisms play a major role in stone biodeterioration. Among unicellular cyanobacteria, Microcystis aeruginosa stands out for its presence in harmful algal blooms (HABs in eutrophic freshwater environments, including urban ponds and artificial water systems, where it can dominate cyanobacterial communities and produce microcystins (hepatotoxic compounds that pose serious risks to aquatic ecosystems and human health) [3,7,8]. Chroococcus sp. is another significant example, generating an exopolysaccharide matrix that promotes biofilm formation and mats, accelerating stone decay [9,10].
Filamentous cyanobacteria such as Nostoc sp., Oscillatoria sp., Phormidium sp., Anabaena sp., and Pseudoanabaena sp. are endolithic and excrete organic acids that dissolve calcium carbonate, weakening the stone and promoting erosion [2,3,11,12]. Biofilm and mat formation intensifies deterioration by creating moist, acidic environments that favor other microorganisms and pollutant accumulation [2,13,14].
Among planktonic green algae, Chlorella vulgaris is frequently involved in biodeterioration. It forms biofilms on monument surfaces and colonizes cracks and pores, trapping moisture and causing thermal expansion–contraction cycles that intensify damage [15]. Filamentous green algae such as Spirogyra sp., Ulothrix sp., Cladophora sp., and Klebsormidium sp. strongly adhere to surfaces, accumulate heavy metals, and promote mechanical fragmentation, further contributing to deterioration [3,12,16,17,18,19].
Traditional algal control methods involve both physical and chemical treatments. Physical approaches include mechanical removal using brushes, scalpels, or spatulas, as well as abrasive air tools, low- or high-pressure water jets, and techniques based on heat shock, ultraviolet (UV-C) irradiation, microwave heating systems, or dry ice treatment. Although these methods avoid the use of chemical agents, they are often insufficient for long-term control and can damage the stone surface by increasing roughness, generating microcracks, or altering texture, thereby enhancing future microbial recolonization. Chemical treatments such as sodium hypochlorite, chlorine tablets, ammonia, or quaternary ammonium biocides can also irreversibly damage stone and harm the ecosystem [19,20,21,22,23,24,25,26]. Chemical treatments such as sodium hypochlorite, chlorine tablets, ammonia, or quaternary ammonium biocides can also irreversibly damage stone and harm the ecosystem. Resistance has been documented in algae such as Chlorella sp., Bracteococcus sp., Scenedesmus sp., Chlorosarcinopsis sp., Pleurocapsa sp., Chroococcus sp., and Chamaesiphon sp. [3,4]. Moreover, recolonization often occurs quickly due to inherent resistance, dormant cell reactivation, biofilm protection, and the bioreceptive nature of stone. Chlorosarcinopsis and Bracteococcus, found in monumental fountains such as those in the Alhambra, show notable tolerance to repeated chemical applications [2,3,27,28].
These limitations have driven interest in natural, non-polluting algicides from plant, microbial, or mineral sources. Such compounds offer sustainable alternatives that reduce environmental and health risks. Studies have tested plant extracts, essential oils, enzymes, and microorganisms for their ability to selectively inhibit algal growth. Integrating these natural approaches into management strategies is a promising avenue for protecting aquatic and stone environments [29,30,31,32,33,34].
Bacteria are already used in cultural heritage conservation, for example, in biocleaning to remove sulfates, nitrates, and black crusts from stone, and to eliminate organic residues such as animal glue and casein from previous restorations [29,30]. They have also been used to remove graffiti [31] and in carbonatogenesis, which promotes calcium carbonate precipitation to consolidate and protect stone [35,36].
However, research on using microorganisms to control algal biodeterioration in stone heritage is scarce [37]. Algicidal bacteria could provide a sustainable alternative to chemical treatments [37,38,39]. This review compiles current knowledge on this approach, identifies research gaps, and explores potential directions for development and practical applications.
This study begins with an overview of the genera and species of microorganisms relevant to controlling biodeterioration in stone heritage. It draws primarily on research on HAB control, while also incorporating examples from other conservation and restoration contexts. We then analyze the mechanisms by which bacteria inhibit algal growth and the composition of the principal algicidal compounds they produce. In addition, we examine the potential for combining algicidal and denitrifying activities to improve sustainability. Finally, we assess the viability of these treatments for real-world heritage applications and discuss the most effective implementation methods. We also compare biological control with traditional methods.
In summary, this paper synthesizes current knowledge on algicidal bacteria, focusing on their potential application to the conservation of stone cultural heritage. It addresses a key research gap by assessing their feasibility as sustainable tools to control algal growth on historic monuments, thus helping to mitigate a major driver of biodeterioration.

2. Algicidal Bacteria and Possible Applications for Heritage Sites

We have compiled and systematized information on 72 algicidal bacteria, creating a database to support future research on their potential use in cultural heritage conservation (Table 1). We selected only bacteria reported to inhibit the freshwater photosynthetic microorganisms most frequently found in fountains and stone monuments affected by biodeterioration. The target includes representants of the main biological groups: cyanobacteria (e.g., Microcystis, Oscillatoria), green algae (e.g., Chlorella, Scenedesmus) and diatoms (e.g., Ulnaria acus).
Most data come from studies on harmful algal blooms (HABs), which occur when algae proliferate excessively and release toxins that disrupt aquatic ecosystems [127], posing risks to aquatic life and human health [128,129]. Although HABs mainly affect natural water bodies, their control strategies can be adapted to manage algal growth in ornamental fountains and other heritage water features, offering potential synergies between aquatic ecosystem protection and cultural heritage preservation. Additional information comes from industrial applications as biogas production and others, where algicidal bacteria are used in co-cultures with microalgae to disrupt cells, enabling the extraction of macromolecules such as lipids and carbohydrates [39,130,131,132,133].
Analysis of the compiled data shows that the most studied photosynthetic microorganism in both HAB and heritage contexts is Microcystis aeruginosa, linked to 56 different algicidal bacteria. Among filamentous cyanobacteria, the most reported genera are Oscillatoria (notably O. tenuis, with 11 associated bacteria), Anabaena (A. flos-aquae, with 7), Nostoc, Pseudanabaena, Phormidium, and Leptolyngbya. For green algae, Chlorella vulgaris (with 9 associated bacteria) is the most studied, followed by Chlamydomonas (6), Scenedesmus, and Choricystis minor. Filamentous green algae such as Spirogyra gracilis and diatoms such as Synedra (Ulnaria) are less frequently recorded.
Several bacterial genera show broad-spectrum algicidal activity against both cyanobacteria and green algae. These include Aeromonas sp., Bacillus sp. (particularly B. cereus), and Pseudomonas sp., outstanding Pseudomonas putida for its activity against multiple strains of Microcystis, Anabaena, and Chlorella. Other versatile strains include Streptomyces lushanensis sp. nov. and Stenotrophomonas sp., which inhibit unicellular and filamentous cyanobacteria, green algae, and even diatoms. This broad activity positions them as promising candidates for biocontrol in heritage water systems, where biofilms often contain mixed microbial communities.
From a phylogenetic perspective, Aeromonas and Pseudomonas belong to the phylum Pseudomonadota (class Gammaproteobacteria), according to the current prokaryotic nomenclature [134], which is the most represented among algicidal bacteria [135], while Bacillus (Bacillota) and Streptomyces (Actinomycetota) also include strains with high inhibitory potential [136,137]. Their ecological adaptability allows them to function under a wide range of environmental conditions [130], making them suitable for sustainable, long-term control strategies in both environmental and heritage contexts.
Several of these genera have already been used in cultural heritage conservation for purposes other than algal control. Examples include Bacillus species with antifungal properties against Fusarium oxysporum, Aspergillus niger, Cladosporium, Penicillium, Mucor, and Alternaria [21,138], and Bacillus subtilis, B. megaterium, and B. pumilus, which have been shown to protect stone surfaces from microbial colonization, though sometimes altering coloration [139,140]. Stenotrophomonas maltophilia has been applied to degrade protein residues from past restorations [141], Ochrobactrum sp. to remove animal glue from paper [142], and extracts from Bacillus flexus and Exiguobacterium undae to clean protein-based residues [143].
Beyond biocleaning, some bacteria contribute to metal stabilization, such as Aeromonas sp. CU5, which transforms corrosion products into stable minerals [144]. In stone bioconsolidation, or carbonatogenesis, strains such as Bacillus subtilis, B. megaterium, Brevibacterium frigoritolerans, B. simplex, B. thuringiensis, B. cereus, Pseudomonas sp., and Myxococcus xanthus promote calcium carbonate precipitation, reinforcing and protecting stone [35,145,146,147]. Other applications include the removal of black crusts from stone and mural paintings using sulfate-reducing bacteria such as Pseudomonas stutzeri [148].
These examples highlight the versatility of bacteria with algicidal potential, demonstrating their ability to combine biofilm control with cleaning and material stabilization. Integrating these functions could enable the development of effective, low-impact, and sustainable biorestoration treatments for cultural heritage in aquatic and humid environments.
Understanding the specific mechanisms by which these bacteria inhibit algal growth is essential for optimizing their application in heritage conservation, ensuring targeted action while minimizing unintended effects on the surrounding environment.

3. How Do Bacteria with an Algicidal Capacity Act?

Algicidal bacteria inhibit algal growth through direct or indirect mechanisms. Direct mechanisms require physical contact with the target algal cells, as observed in Pseudomonas fluorescens, which specifically adheres to diatoms and induces cell lysis [57]. Similarly, Cytophaga sp., Paucibacter aquatile DH15, Stenotrophomonas sp. KT48, and various Bacillus species eliminate Microcystis aeruginosa via this approach [37,71,78,131]. Cell lysis is among the most studied processes due to its high efficiency in reducing algal biomass, offering a safe alternative for human health, the environment, and the preservation of cultural heritage [136].
Indirect mechanisms involve the release of bioactive compounds such as antibiotics, biosurfactants, amino acids, alkaloids, or enzymes, which inhibit algal growth without direct contact (Figure 3) [75,107,149]. These compounds can disrupt key algal metabolic processes, for example, by impairing photosystems I and II during photosynthesis, or by inducing oxidative stress through enzyme inactivation and disturbance of intracellular ion homeostasis [49,57,75].
Studies conducted between 2016 and 2021 identified indirect action as the most common mechanism among algicidal bacteria [37,128,150]. Combining indirect with direct mechanisms enhances overall effectiveness [78]. Additionally, reducing essential nutrients such as nitrogen and phosphorus further restricts algal proliferation in eutrophic waters, improving the performance of algicidal bacteria [74].

4. Main Metabolites with Algicidal Capacity Excreted by Bacteria

According to Meyer et al. (2017) [129], enzymes secreted by algicidal bacteria constitute one of the main indirect mechanisms of action. They inhibit algal growth by degrading essential structural and functional cell components. Proteases and lipases from Flammeovirga yaeyamensis hydrolyze cell wall proteins and membrane lipids, compromising membrane integrity and leading to cell lysis or leakage of intracellular contents, thereby suppressing the growth of Chlorella vulgaris [119]. L-amino acid oxidase (L-AAO), secreted by Aquimarina sp., catalyzes the oxidative deamination of amino acids, generating hydrogen peroxide that induces oxidative stress and damages organelles and photosynthetic pigments, resulting in algicidal activity against Microcystis aeruginosa, Phormidium persicinum, Chlamydomonas raudensis and Chlorella pyrenoidosa [114].
A second major group of algicidal compounds comprises amino acid derivatives, peptides, and protein-like molecules, particularly cyclic diketopiperazines (DKPs). These stable dipeptides are synthesized by genera such as Bacillus, Stenotrophomonas, Chryseobacterium, and Aeromonas [65,92,151,152,153]. For example, Bacillus sp. Lzh-5 produces cyclo(Gly-Pro) and cyclo(Pro-Val) [66]. Stenotrophomonas sp. F6 also produces cyclo(Gly-Pro) along with hydroquinone, a phenolic compound with broad algicidal activity [92]. Chryseobacterium sp. GLY-1106 synthesizes two DKPs: 1106-A cyclo(4-OH-Pro–Leu) and 1106-B cyclo(Pro–Leu) active against various cyanobacteria and green algae, including Microcystis aeruginosa, M. viridis, Oscillatoria sp., Synechococcus sp., and Chlamydomonas sp. [50]. Within this group, Aeromonas sp. GLY-2107 also stands out for producing two potent algicidal compounds: 3-benzyl-piperazine-2,5-dione (a DKP) and 3-methylindole (an indole derivative), both effective against M. aeruginosa, M. viridis, Chroococcus sp., Oscillatoria sp., and Chlamydomonas sp. [53].
This category also includes aromatic and indole-derived compounds, many of which are associated with significant algicidal activity. Aeromonas veronii produces lumichrome, a riboflavin catabolite with confirmed activity against M. aeruginosa [48]. Similarly, the indole alkaloid tryptoline, produced by Streptomyces eurocidicus and Bacillus siamensis, has demonstrated inhibitory activity against several algal species [65]. Other tryptophan-based compounds, such as tryptamine, and pigments like prodigiosin, secreted by Serratia marcescens, Enterobacter hormaechei, and Hahella sp., also display strong algicidal effects [72,89,154,155].
Additional nitrogenous metabolites, such as bacilysin and various purine derivatives, are produced by several Bacillus species with demonstrated algicidal activity. Bacillus amyloliquefaciens FZB42 synthesizes bacilysin [58], while purine derivatives have been identified in Bacillus thuringiensis Q1 [70], all showing inhibitory effects on Microcystis aeruginosa, Anabaena flos-aquae, Chlorella vulgaris, and Oscillatoria sp.
Within the group of organic acids and related compounds, Raoultella ornithinolytica has been reported to secrete metabolites such as D-gluconic acid, chlorogenic acid, L-malic acid, 5-hydroxy-2,4-dioxopentanoate, and 2-methyl-3-oxopropanoic acid, all contributing to its inhibitory effect on M. aeruginosa [84].
Finally, some bacteria exert algicidal activity through flocculation-based mechanisms, promoting aggregation and sedimentation of algal cells. Paebubacillus sp. A9 produces carboxylated polysaccharides that act as natural flocculants, effectively reducing microalgal abundance in aquatic systems [38].
These findings, summarized in Table 2, highlight the structural and functional diversity of bacterial algicidal metabolites. Their biological origin and versatility support their use as sustainable alternatives to conventional biocides, particularly in sensitive environments and heritage contexts. Although mechanisms vary, many act by inducing oxidative stress, disrupting cell membranes, or inhibiting essential metabolic pathways. Among the classes reviewed, diketopiperazines and low-molecular-weight peptides such as bacilysin stand out for their chemical stability, positioning them as promising tools for ecological and responsible algal bloom control.

5. Denitrifying Algicidal Bacteria: A Combined Approach for Stone Cultural Heritage Conservation

Salt efflorescence is a frequent and damaging alteration in stone heritage, manifesting as whitish deposits on the surface of construction materials. It occurs when soluble salts dissolved within the stone migrate to the surface and crystallize as water evaporates. Repeated crystallization leads to loss of material cohesion, causing cracks, detachments, and even disintegration of the stone, ultimately compromising both the structural integrity and esthetics of historical monuments [163,164].
Nitrates are among the most problematic salts in this process. In addition to contributing to salt crystallization, they increase the bioreceptivity of stone surfaces by serving as a nutrient source for phototrophic microorganisms. This dual role makes nitrate control particularly relevant for preventing both biodeterioration and salt-related mechanical damage in heritage contexts [165,166,167].
In this regard, bacteria with both algicidal and denitrifying capabilities represent an innovative and sustainable strategy for heritage conservation. A notable example is Streptomyces sp. L1, which not only eliminates algae by inhibiting photosynthesis (reducing up to 85% of Microcystis aeruginosa) but also significantly decreases nitrogen levels in water, including nitrates, nitrites, and ammonium [95]. This dual functionality is critical for controlling harmful algal blooms and preventing eutrophication, while also mitigating the formation of salt efflorescence, thereby addressing two key factors in the deterioration of cultural assets [168].
Other bacteria, such as Pseudomonas stutzeri [169] and Bacillus sp. [170], have also been studied for their denitrifying capacity in conservation and restoration treatments. Recent research has highlighted their algicidal activity, making them promising candidates for integrated biocleaning strategies [140,163,169,171]. Incorporating these bacteria into conservation protocols could enhance the effectiveness and environmental compatibility of treatments, supporting a holistic and sustainable approach to heritage preservation.

6. Synergies Among Bacteria to Control Algal Growth

Algicidal bacterial consortia offer an effective strategy to inhibit algal growth by means of different microorganisms acting in synergy. This approach is based on the complementary action mechanisms of each strain, allowing for broader and more efficient control of harmful algae [128,136]. Synergistic interactions among strains often lead to more persistent and reliable algal inhibition, especially in natural and variable environments where individual strains may lose efficacy over time. These findings, mostly derived from HAB control studies, provide a transferable framework for heritage contexts.
He et al. (2021) [49] explored the use of multispecies algicidal combinations as a sustainable strategy to control harmful algal blooms in freshwater. Using coconut fiber carriers, bacterial communities were enriched which significantly inhibited the growth of various algae, including cyanophytes, diatoms, and dinoflagellates. Genera like Bacillus and Pseudomonas were noteworthy for their ability to release substances that damage algal cell membranes. This system showed an efficiency of 53.74%, underlining the stability and effectiveness of multispecies communities compared to individual strains.
Another example of this synergy is the consortium composed of Rhizobium sp., Methylobacterium zatmanii, and Sandaracinobacter sibiricus, which achieved a 95% lysis of Microcystis aeruginosa cells in 10 days, significantly surpassing the results obtained by each species individually (75–82% lysis). This consortium not only inhibits the growth of M. aeruginosa but also degrades microcystins, the toxins produced by these cyanobacteria [85].
Synergy can also enhance the spectrum of activity. While bacteria often act selectively against certain algal taxa, their combination can expand their range of inhibition. For instance, the consortium of genera Bacillus and Aeromonas has been reported to eliminate Chroococcus [49]. This illustrates the potential of multispecies bacterial consortia to act simultaneously against diverse phototrophic groups, including cyanobacteria, diatoms, and green algae, thereby broadening their ecological effectiveness and adaptability.
A particularly illustrative case of synergy and complementarity is the dual action of Aeromonas bestiarum HYD0802-MK36 and Pseudomonas syringae KACC10292T against M. aeruginosa. While HYD0802-MK36 exerts a direct algicidal effect through physical contact with cyanobacterial cells, KACC10292T acts indirectly by releasing algicidal compounds into the medium. Notably, the algicidal activity of P. syringae increased when co-cultivated with its algal target, suggesting that interaction may stimulate a greater production of inhibitory compounds. Together, these strains illustrate how combining direct and indirect algicidal mechanisms can enhance efficacy and adaptability across diverse environmental conditions [56].
In order to ensure the stability and efficacy of these treatments, it is essential to select biocompatible bacteria that, whenever possible, do not compete with each other for the same resources. Despite notable progress, further research is needed to identify optimal combinations of algicidal bacteria adapted to different algal species and environmental conditions, particularly in complex systems such as ornamental water fountains or cultural heritage sites. These principles, well established in HAB management, could guide the design of consortia tailored for heritage conservation, a topic further developed in the Discussion section.
The synergistic use of algicidal bacteria represents a promising and sustainable approach to control phototrophic growth in heritage fountains. As previously discussed, multi-strain consortia are generally more stable and adaptable than single strains, providing longer-lasting inhibition of algae and cyanobacteria in dynamic aquatic systems. However, these cooperative interactions must be carefully optimized, as some combinations may unintentionally stimulate non-target microorganisms. Therefore, controlled testing and case-specific formulations are essential before applying multispecies systems to heritage environments.

7. Methods in Practice

The methods for applying bacteria to control harmful algal blooms (HABs) cover a variety of strategies. Coyne, Wang, and Johnson (2022) [128] provide an in-depth review of approaches to using algicidal bacteria in managing HABs:
1. Direct dispersion of bacteria or algicidal products: This approach involves releasing live bacteria or their bioactive compounds directly into aquatic environments [7,39]. For example, Xanthobacter autotrophicus was applied in situ in a small artificial concrete pond connected to the Han River Reservoir in Korea to control Microcystis aeruginosa. Algicidal activity was observed, but periodic inoculations were necessary to maintain effectiveness [100]. This method faces several limitations, including degradation of algicidal compounds, consumption by other aquatic organisms, dispersion by water currents, and potential adverse effects on biodiversity and non-target organisms [7].
2. Immobilized bacteria: In this method, bacterial cells are encapsulated or fixed within carriers that allow controlled, localized release of bacteria or their algicidal compounds [137]. Common carriers include polyvinyl alcohol–sodium alginate (PVA) [172], often with additives such as Fe3O4 and wheat bran [41,173]; alginate hydrogel [174]; activated carbon-PVA cellulose sponge [175]; and various natural materials such as polyurethane, polyester, agarose, and coconut fiber agar [176].
3. Multifunctional systems: These involve the co-immobilization of several algicidal strains to achieve simultaneous or complementary effects against HABs [38,177]. This method has been applied in wastewater treatment [178] and in eliminating harmful algae and their associated toxins released during bacteria–algae interactions [179]. Another promising approach is the development of genetically engineered algicidal bacteria with dual functions [180,181]. However, research on multifunctional systems for HAB mitigation and prevention remains limited [39]. Coyne, Wang, and Johnson (2022) suggest exploring strategies combining immobilized non-harmful algae with specific bacteria to reduce nutrient levels while directly inhibiting HAB species [128].
4. Recruitment of natural algicidal bacteria: Certain substrates, such as seagrass meadows or macroalgae, can attract and concentrate native algicidal bacteria [39,176,182]. This low-impact approach leverages natural ecological interactions to control algae in sensitive areas, minimizing environmental disturbance [183,184].
For biocleaning in cultural heritage assets, the methods can be adapted from those used in HAB control. Variations in these techniques have already been implemented in conservation and restoration treatments on stone materials:
1.
Direct application:
Immersion of heritage materials in bacterial cultures has shown some efficacy. For example, treatments using Desulfovibrio desulfuricans on marble and sandstone partially removed sulphated crusts and alteration layers within 60–84 h [185,186,187,188,189,190]. However, this method is impractical for large or fragile objects, and post-treatment drying can cause salt migration and efflorescence, as well as mechanical stresses from uneven water retention, leading to cracking, exfoliation, or fractures. The presence of embedded metallic elements can also promote corrosion during prolonged moisture exposure [164,187,191].
To overcome these drawbacks, spray application offers a practical alternative, particularly for complex geometries or surfaces requiring minimal contact. While sprays allow rapid, uniform coverage, reduced contact time can lower efficacy unless combined with moisture-retaining wraps or gels. Furthermore, this method poses a higher risk of uncontrolled environmental dispersion, requiring strict control of application areas and conditions [21].
More recently, an innovative variant of direct application, known as dry biocleaning, has been introduced. This method involves the use of lyophilized microorganisms or dehydrated cells directly on the stone surface. A representative case is the dry biocleaning system with Saccharomyces cerevisiae, in which yeast cells are mixed with powdered sucrose and manually applied onto the substrate. Ambient humidity reactivates the cells, triggering fermentative metabolism that facilitates the removal of soluble salts and atmospheric deposits within 12-18 h. This procedure, tested on travertine from the Quattro Fontane fountain in Rome, proved effective in reducing nitrates, sulfates, and other contaminants, producing a significant chromatic improvement without adverse effects detected during short- and medium-term monitoring [192].
2.
Immobilization:
Specific carriers are used for controlled release of bacteria and their algicidal compounds. Each and every material has specific characteristics that make it suitable for different types of cleaning. Bosch-Roig, Lustrato, Zanardini and Ranalli (2015) and Ranalli and Zanardini (2021) [164] review the types of system delivery for biocleaning treatment on cultural heritage elements.
For example, Laponite®RD (CTS Spain, Madrid, Spain), a synthetic clay composed of silicates, forms a gel that allows it to be used in the restoration of cultural heritage [193]. This type of carrier has the advantage that it can be used on vertical surfaces [189]. It has been used with Cellulosimicrobium cellulans TBF11E to remove inorganic sulfate deposits, Stenotrophomonas maltophilia UI3E for organic substrates of protein origin, and Pseudomonas koreensis UT30E for phosphates and various organic materials. This method was used to restore the wall paintings in the lower loggia of the Casina Farnese in Rome [141].
Carriers such as cotton wool and cellulose pulp (Arbocel®) (CTS Spain, Madrid, Spain) are inexpensive and easy to use, and do not interfere with cellular activity [187,194]. Nevertheless, they may be less effective than Carbogel® (CTS Spain, Madrid, Spain) or Laponite® (CTS Spain, Madrid, Spain) for applications requiring greater stability or adhesion, especially at low concentrations [189].
Sepiolite has been used for biocleaning on stone with desulphating bacteria such as Desulfovibrio vulgaris [170]. These bacteria act in anaerobic conditions to eliminate the sulfates present in the stone, using sepiolite as a carrier to maintain humidity and foster biological activity without the presence of oxygen [193,195]. They have also been used to remove black crusts [196]. However, they have the potential to release traces of iron, which can darken the treated surface [189].
Hydrobiogel 97, used with D. vulgaris to remove black crusts on Failaka Island (Kuwait), maintains the humidity necessary for bacterial activity, but may be less effective in treatments requiring prolonged applications [197]. In addition, it is difficult to remove it completely after treatment due to its strong adhesion. Applying it on non-horizontal surfaces can become difficult due to its excessive fluidity, which restricts its use in certain cases [164].
Agar has good water retention and is easy to apply on flat surfaces, but has low adhesion on rough surfaces. Even so, it is suitable for biocleaning organic waste and salts [164,189]. Mortar with alginate beads, used on sandstone walls in Matera (Italy) to remove nitrates with Pseudomonas pseudoalcaligenes and D. vulgaris, provides a suitable medium for the controlled release of microorganisms and effective treatment of saline deposits [164,198]. It provides precise dosages [189], though its high adhesiveness and difficulty to remove it makes its use inadvisable, due to the possible damage it may cause on the surfaces of cultural assets.
Carbogel® (CTS Spain, Madrid, Spain) is mainly used in the removal of black crusts and sulfate deposits on sculptures and stone surfaces such as marble and limestone. This material enables a microaerophilic condition to be created when covered with a thin film, which fosters biological activity in repeated 15 h treatments, achieving up to 98% sulfate removal in specific tests [170]. It has been used to restore sculptures such as Michelangelo’s Rondanini Pietà [196,199]. This is noteworthy for the ease in applying and removing it, and its high bacterial compatibility. However, it has limitations as regards adhesion and can be difficult to handle due to its less compact structure, especially on vertical surfaces [164,189].
3.
Recruitment of bacteria in natural environments:
Carbonatogenesis uses indigenous stone-inhabiting bacteria for bioconsolidation. Specific nutrient solutions stimulate native populations to precipitate calcite, strengthening the substrate [200,201]. This method has been applied at the Monastery of San Jerónimo (Granada, Spain), successfully enhancing cohesion through the activity of native carbonatogenic bacteria [202]. While widely studied for structural consolidation, its application to removing biofilms or algae has not yet been documented.

8. Algicidal Bacteria Versus Traditional Biocides: Advantages and Drawbacks

Conventional methods for controlling microalgal growth in heritage conservation, including chemical biocides and mechanical techniques, have significant limitations in terms of efficacy, material safety, and environmental impact. Chemical treatments, particularly those based on quaternary ammonium compounds (QACs) such as benzalkonium chloride (BAC), didecyldimethylammonium chloride (DDAC), and other nitrogen- or phenol-derived agents, target a broad spectrum of microorganisms [74,203]. While they can be effective in the short term, repeated applications may alter the physicochemical properties of treated substrates and promote algal resistance [27,28,204,205,206]. This necessitates frequent reapplications, increasing the risk of cumulative material damage (Figure 4). Moreover, compounds like BAC-C12 have been shown to induce microcystin release from Microcystis aeruginosa, thereby exacerbating environmental toxicity [207,208].
Mechanical cleaning methods, although widely used, rely on abrasive tools that can compromise the structural integrity of heritage surfaces. These techniques generally act only on the surface and are ineffective against endolithic algae that penetrate deeply into the stone matrix. The invasive nature of such interventions limits their long-term viability, particularly when repeated treatments are required [2,28].
In contrast, algicidal bacteria can provide a more sustainable and selective alternative. Their specificity enables targeted inhibition of harmful algal species without affecting beneficial or neutral microorganisms or damaging the substrate [39,155,209]. For example, Bacillus flexus secretes harmine and norharmane to inhibit the growth of cyanobacteria, without affecting other phytoplankton species [209]. Similarly, the cell-free filtrate of Streptomyces sp. LJH-12-1 (L1) strongly inhibited M. aeruginosa while having no impact on green algae such as Chlorella vulgaris and Scenedesmus obliquus, indicating selective activity toward cyanobacteria [95]. These bacteria are cost-effective to culture, environmentally compatible, and non-aggressive to historical materials [210]. Furthermore, their metabolites are generally rapidly biodegradable, reducing the risk of long-term toxicity and environmental accumulation [30,155,211].
Unlike chemical biocides, biological treatments can incorporate native microbiota in processes such as bioconsolidation. However, previous treatments with broad-spectrum chemical biocides may disrupt or eliminate these communities, necessitating the introduction of exogenous strains [36]. This underscores the ecological value of biological control agents, particularly when the aim is to conserve or restore the microbial balance of heritage surfaces, as shown in studies on both stone and metal substrates [144,212]. Although individual strains may exhibit limited action spectra, broader efficacy can be achieved through bacterial consortia or synergistic interactions. Nevertheless, these approaches remain underexplored and require further evaluation under real-world conditions [28].
In addition to bacterial compounds, recent research has highlighted the potential of microbial metabolites as natural biocides which can be used in combined treatments. For example, Petraretti et al. (2022) [213] evaluated four fungal secondary metabolites (cavoxin, epi-epoformin, seiridin, and sphaeropsidone) and found them effective against biodeteriogenic fungi such as Aspergillus niger, Fusarium oxysporum, and Alternaria alternata, with significantly lower ecotoxicity than commercial biocides like Preventol® (CTS Spain, Madrid, Spain) and Rocima® (CTS Spain, Madrid, Spain).
Moreover, the use of cell-free filtrates (bioactive substances secreted by bacteria without the introduction of living organisms) offers a promising option when the direct application of microorganisms may pose ecological or material risks. These compounds combine the selectivity and biodegradability of biological systems with greater control and safety in sensitive conservation contexts [57,117,154,207].
Overall, algicidal bacteria represent a biologically based and environmentally responsible strategy for heritage conservation. While their application still presents challenges, such as longer treatment durations and the need to monitor potential microbial proliferation, their potential as an alternative to traditional biocides is considerable and warrants continued interdisciplinary research [46,49,214].

9. Discussion

9.1. Bacteria for Controlling the Growth of Photosynthetic Microorganisms in Stone Heritage

As explained throughout the article, the current understanding of bacteria with algicidal potential is primarily based on studies focused on harmful algal blooms (HABs), especially in aquatic environments. However, the possible applications of these microorganisms for the control of many microalgae and cyanobacteria frequently involved in the biodeterioration of ornamental fountains and stone heritage remain largely unexplored. Filamentous cyanobacteria such as Leptolyngbya, Calothrix, and Schizothrix, along with unicellular forms like Pleurocapsa, Gloeocapsa, Chroococcidiopsis, and Chamaesiphon, are recurrent in stone surfaceses [2,3] but have not been the focus of bacterial biocontrol studies. The same applies to filamentous green algae such as Klebsormidium, Stigeoclonium, Ulothrix, and Cladophora, and coccoid green algae from genera like Bracteococcus, Stichococcus, Chlorosarcina, and Apatococcus, which are widely documented in biodeterioration processess [12,14] yet remain understudied from a microbiological control perspective.
Freshwater diatoms also represent a neglected group in this field. Most algicidal research on diatoms focuses on Stephanodiscus hantzschii in HABs [215,216], a species not typically involved in monument colonization. Nevertheless, a study by Zakharova et al. (2013) [126] demonstrated that certain bacteria can degrade diatom cells and contribute to silicon recycling. This opens a promising research avenue for identifying bacteria capable of targeting diatom genera commonly detected in stone biodeterioration, such as Navicula, Nitzschia, Achnanthes, and Melosira, among others [2,3,12,14,16].
Despite these gaps, several bacterial genera already studied in HAB research show clear translational potential for heritage conservation, including Achromobacter, Acinetobacter, Aeromonas, Chryseobacterium, Flavobacterium, Ochrobactrum, Paucibacter, Paenibacillus, Pseudomonas, Stenotrophomonas, Streptomyces and Bacillus (Table 1). These encompass broad-spectrum algicidal activity and traits compatible with stone substrates, positioning them as strong candidates for adapted conservation protocols.
The genus Bacillus, which includes well-studied species such as B. cereus and B. subtilis, is the most extensively characterized for the control of HABs, acting against a wide spectrum of photosynthetic microorganisms. Importantly, some Bacillus strains are already integrated into heritage conservation practices, confirming their safety and compatibility with sensitive materials [30,35,217,218,219]. Their successful use in different restoration contexts supports the feasibility of extending Bacillus-based strategies to the biocontrol of phototrophic colonization in stone monuments.
However, there are other genera and bacterial strains with great potential to be incorporated into conservation-restoration treatments aimed at controlling the growth of microalgae. In particular, bacteria from the phylum Actinomycetota represent a significant opportunity, due to their functional versatility, low pathogenicity, and environmental compatibility. Several strains within this order have demonstrated inhibitory activity against both cyanobacteria and green algae, broadening their applicability for diverse biodeterioration scenarios [95,99]. Although their use in cultural heritage conservation has not yet been systematically explored, their properties position them as promising candidates for respectful interventions involving stone materials and surrounding ecosystems.
Within this group, species of the genus Streptomyces, such as S. globisporus and S. lushanensis [93,96], have shown notable allelopathic effects against a broad spectrum of microalgae involved in biodeterioration processes. Additionally, the strain Streptomyces sp. L1 has proven effective in inhibiting the growth of Microcystis aeruginosa and significantly reducing dissolved nitrogen levels in the water (nitrates, nitrites, and ammonium) [95]. Furthermore, siderophores produced by Streptomyces spp., such as desferrioxamines G, B, and E, have been successfully applied in the conservation of cultural heritage due to their high specificity in chelating iron. These hydroxamate compounds enable the selective removal of ferric stains from lignocellulosic supports, textiles, and paper, without altering the integrity of the original material. Their use represents a sustainable biotechnological alternative to conventional chemical methods, minimizing substrate damage [140].
On the other hand, strains such as Paucibacter aquatile DH15 [78], Paebubacillus sp. A9 [38], Acinetobacter sp. J25 [103], Bacillus subtilis S4 [69] and Achromobacter xylosoxidans have the capability to degrade microcystins, toxins produced by M. aeruginosa, improving water quality [43]. These bacteria could also be relevant for targeting other cyanobacterial genera associated with microcystin production and stone biodeterioration, such as Anabaena, Pseudanabaena, Nostoc, and Oscillatoria, contributing not only to the preservation of submerged cultural heritage but also to broader environmental remediation strategies.

9.2. Selectivity in Treatments

Bacteria offer a valuable opportunity for designing species-selective treatments, as some strains inhibit specific algal targets while leaving other photosynthetic taxa unaffected [57,207]. For instance, certain Paucibacter strains can suppress M. aeruginosa without impacting non-target cyanobacteria or green algae [78].
Likewise, Bacillus thuringiensis Q1 [70] and Bacillus flexus SSZ01 [108] selectively inhibit cyanobacteria while sparing green algae. Bacillus sp. SY-1, although a marine bacterium, produces micosubtilin MS 1084, which is active against dinoflagellates and raphidophytes but shows little effect on both marine and freshwater microalgae [137].
These findings highlight the species-specific nature of algicidal compounds, a feature desirable for targeted control. However, further studies are needed to assess their ecological impact on microbial communities and higher trophic levels [183,207]. While selectivity minimizes collateral effects, it may limit efficacy in multispecies algal biofilms, where bacterial consortia may provide broader and more robust control.

9.3. Synergies Between Bacteria

The synergistic application of algicidal bacteria represents a promising and sustainable approach for the long-term control of phototrophic growth in ornamental and heritage water systems. As demonstrated by recent studies, multi-strain consortia can provide enhanced stability, adaptability, and algicidal performance compared to single-strain treatments [49]. Commercial formulations based on Bacillus consortia further illustrate the feasibility of this strategy. In aquaculture, these probiotics have been employed to improve overall water quality by reducing nitrogenous compounds, phosphates, organic matter, and pathogenic microbes. Products such as Sanolife PRO-F® (B. subtilis, B. licheniformis, and B. pumilus) (INVE Aquaculture, Dendermonde, Belgium) [220], EcoAqua (B. subtilis, B. licheniformis, B. megaterium, and B. laterosporous) (EcoMicrobials LLC, Miami, Florida) [221], and Efinol® L (B. subtilis, B. licheniformis, Lactobacillus acidophilus, and Saccharomyces cerevisiae) (CODEMET S.A., Guayaquil, Ecuador) have proven effective in regulating key water parameters under intensive conditions [222]. Notably, Ecotrax® (a mixture of seven Bacillus species) (TIL Biosciences, Chennai, Tamil Nadu, India) enhanced water transparency by limiting microalgal proliferation while simultaneously improving physicochemical stability [223]. These examples highlight that multi-strain Bacillus formulations can act as effective bioremediation tools, with direct relevance to ornamental ponds in historical gardens, where algal colonization represents a recurrent conservation challenge. The application of bacilli in bioremediation is inexpensive, straightforward, and environmentally sustainable [224], which further supports its consideration for cultural heritage contexts.
This strategy has already been explored in the heritage field. A microbial consortium composed of Bacillus cereus, Bacillus subtilis, and Exiguobacterium sp. was used to clean marble surfaces of the sculpture La Lupa by G. Graziosi at the National Gallery of Modern and Contemporary Art in Rome, achieving the removal of urban pollution deposits without affecting the original material [139]. These results highlight the potential of biobased interventions as an alternative to conventional chemical treatments, particularly for complex substrates such as stone or marble.
Nevertheless, certain limitations must be considered before implementing multispecies bacterial treatments. Some strains, despite showing broad algicidal activity, can stimulate the growth of non-target phototrophs, potentially causing new forms of biodeterioration or ecological imbalance. For instance, Pseudomonas fluorescens MZ007859 inhibits various cyanobacteria but promotes Chlorella vulgaris growth [106]; Bacillus amyloliquefaciens T1 suppresses Microcystis aeruginosa yet stimulates Chlorella pyrenoidosa [59] and Pseudomonas putida HYK0203-SK02 inhibits M. aeruginosa and Anabaena cylindrica but enhances the proliferation of the diatom Cyclotella sp. [115]. These cases reveal the complexity of microbial interactions and the necessity of pre-evaluating bacterial compatibility and selectivity to avoid unintended side effects.
In ornamental fountains, photosynthetic microorganisms occur mainly as planktonic species, floating on the water surface, or as benthonic forms attached to stone surfaces or embedded within biofilms [12]. The latter pose a greater conservation challenge because their adhesion (mediated by an extracellular polysaccharide matrix) provides mechanical stability, promotes mineral dissolution, and increases stone roughness, enhancing future bioreceptivity [139]. Physical removal methods such as brushing, pressurized water, or abrasive cleaning are less effective against benthonic phototrophs, which often form compact biofilms or colonize stone pores and microfissures [20,205]. Certain cyanobacteria, such as Pleurocapsa sp. and Chamaesiphon sp., exhibit chasmoendolithic growth, penetrating deep into cracks and fissures where physical methods cannot reach without risking erosion or material loss [2,3,11]. In contrast, algicidal bacteria offer a selective and eco-compatible solution: some strains can degrade the extracellular polymeric substances (EPS) of phototrophic biofilms [225,226,227], weakening their cohesion and facilitating gentle detachment without damaging the substrate. Moreover, being biological systems, they can infiltrate stone pores more naturally, avoiding the risks associated with chemical biocides, which often increase porosity and material sensitivity. Although no adverse effects on stone substrates have been reported so far after the application of bacteria or their metabolites, further research is needed to evaluate potential long-term impacts, particularly in sensitive cultural heritage materials.
In this context, bacterial synergies could provide an effective route to address the challenge of inhibiting phototrophic microorganisms with distinct ecological traits (planktonic, benthonic, or endolithic) that deteriorate stone through different mechanisms. Tailored combinations of algicidal bacteria could maximize biocontrol efficacy while ensuring material compatibility and environmental sustainability.

9.4. Can Microorganisms Involved in Biodeterioration Processes Be Used for the Conservation of Cultural Assets?

Using microorganisms traditionally linked to biodeterioration as conservation tools represents a paradigm shift in heritage management. This approach demands strain-level characterization, as members of the same genus can display opposite effects on materials.
A clear example is the genus Shewanella: while S. loihica has been linked to metal stabilization processes [228,229], S. oneidensis is capable of destabilizing and dissolving the passivating layer of magnetite (Fe3O4), a natural protection against metal corrosion [230]. Similarly, the case of Aeromonas sp. clearly illustrates this duality. This bacterium is common in aquatic environments and has been identified in organic heritage materials such as easel paintings, documents, photographic supports, and gelatin-based adhesives. Its ability to degrade collagen makes it a potential biodeteriogenic agent [29,198,231]. However, certain environmental strains, such as CA23 and CU5, have shown potential for stabilizing corroded metal objects by transforming ferric corrosion products (goethite and lepidocrocite) into more stable ferrous minerals like vivianite and siderite [144]. In the context of microalgal biocontrol, this degradative activity can be advantageous, as Aeromonas sp. feeds on the exopolysaccharides (EPS) present in biofilms [48], facilitating their removal from stone surfaces without the need for chemical intervention.
Selection must consider metabolic traits, the asset type, material composition and intervention goals. Application should be case-specific, supported by applied studies that validate both efficacy and safety under real conservation conditions, and preceded by biosafety screening to avoid introducing opportunistic pathogens.

9.5. The Use of Living Organisms: An Environmental Risk or a Biotechnological Opportunity?

A key concern in applying algicidal bacteria is the ecological risk of releasing living strains. However, many can be used indirectly via the bioactive compounds they secrete, which target HAB-causing microalgae while avoiding direct introduction of microorganisms into aquatic or heritage environments [232].
These compounds have shown low toxicity toward non-target organisms, including fish and aquatic invertebrates [155], and maintain their activity across a wide range of pH and temperature conditions [233,234], enhancing their applicability in real-world settings.
The genus Bacillus combines a strong biosafety record with metabolic diversity, producing cyclic peptides, nitrogenous bases, dipeptides, indole alkaloids, and lipopeptides (surfactin, iturin, fengycin) with algicidal potential [235,236,237]. While often studied for antifungal effects, many of these compounds remain underexplored for algal control. Bacillus is also well-established in conservation (bioconsolidation, biocleaning, salt removal), making it a strategic resource at the interface of HAB mitigation and heritage preservation.
Moreover, numerous studies indicate that cell-free bacterial filtrates are often more effective than living cells, underscoring the importance of further characterizing these compounds and evaluating their behavior under diverse environmental conditions [60,124,150,159]. These filtrates combine the selectivity and biodegradability of biological treatments with greater safety in sensitive heritage environments, as they avoid the introduction of exogenous microorganisms while still enabling targeted control of algal species.
Before conservation use, these compounds require evaluation for stability, efficacy, and ecological impact, including resistance to UV, temperature shifts, moisture variation, and pH changes (conditions typical of outdoor heritage sites). Despite promising results, their long-term performance and environmental interactions remain underexplored, warranting further applied research under real conservation conditions.

10. Conclusions

This review highlights the growing potential of biological methods in the conservation and restoration of cultural heritage, particularly for controlling microalgal growth on stone surfaces. The use of bacteria with algicidal capabilities and their bioactive compounds offers a sustainable, low-impact alternative to conventional physicochemical treatments. These approaches provide greater specificity, deeper penetration, and leave no contaminant residues, in line with conservation principles that emphasize material compatibility and ecological responsibility.
Safe and effective application requires a detailed understanding of strain behavior, metabolite properties, and substrate characteristics. Since effects may vary even within the same genus, and substrate type strongly influences performance, interventions must be tailored to each case and supported by applied research and in situ testing.
Within the wide diversity of reported algicidal bacteria, Bacillus and Streptomyces emerge as the most promising for heritage applications. They combine a strong biosafety record with the production of stable metabolites such as lipopeptides, diketopiperazines, and amino acid derivatives, and have already been successfully employed in biocleaning and bioconsolidation.
Other genera, including Pseudomonas, Aeromonas, and Stenotrophomonas, encompass opportunistic pathogens; however, they exhibit a broad algicidal spectrum, effectively inhibiting both cyanobacteria and green algae. In heritage contexts, the use of their cell-free filtrates or purified metabolites represents a safer alternative, ensuring efficacy while minimizing ecological and health risks, and offering greater control in sensitive environments.
Complementary genera such as Paenibacillus, Acinetobacter, Exiguobacterium, Raoultella, Ochrobactrum, Chryseobacterium, Flavobacterium, and Paucibacter further expand the potential of microbial-based treatments through additional functions, including denitrification, flocculation, and protein degradation.
It is important to note that, while most studies on algicidal bacteria have focused on genera typical of harmful algal blooms (HABs), other phototrophs frequently associated with cultural heritage biodeterioration, such as green algae (Chlorococcum, Haematococcus, Apatococcus, Ulothrix), cyanobacteria (Gloeocapsa, Calothrix, Pleurocapsa, Schizothrix), and diatoms (Navicula, Nitzschia, Achnanthes, Melosira), remain almost completely unexplored. Translating the knowledge generated in the context of HABs to these biodeteriogenic organisms therefore represents a crucial opportunity to design more specific and effective conservation strategies.
Future research should prioritize the development of standardized protocols to evaluate microbial strains and bioactive compounds under realistic conservation conditions, promote interdisciplinary collaboration between microbiologists and conservators, and establish regulatory frameworks that ensure safety and efficacy. These steps are essential to translate laboratory findings into field-ready solutions compatible with heritage requirements.
Microorganisms, once regarded solely as agents of deterioration, are increasingly recognized as valuable allies in heritage preservation. When applied with scientific rigor and environmental responsibility, they open new avenues for integrated, sustainable, and forward-looking conservation.

Author Contributions

Conceptualisation, writing and work design, I.C.-B.; original draft preparation review, supervision and editing, J.R.-N.; project administration, funding acquisition, J.R.-N. and F.B.-G. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the following projects: ECODIGICOLOR, grant number TED2021-132023B-I00, supported by MCIN/AEI/10.13039/501100011033 and Unión Europea NextGenerationEU/PRTR (Proyectos estratégicos orientados a la transición ecológica y digital), project BIOALHAMBRA, grant number PID2022-143064OB-I00, supported by MCIN/AEI/10.13039/501100011033 «Proyectos de Generación de Conocimiento», and project SOL2024-30779, (VII Plan Propio de Investigación-Programa al estímulo de áreas con necesidades investigadoras y de la actividad investigadora emergente—Modalidad A1—by University of Seville).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Data sharing is not applicable to this article.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Biopatina on ornamental fountains: (a). Fountain located in the Royal Alcazar of Seville. (b) Fountain located in the Alhambra and Generalife monument (Granada, Spain).
Figure 1. Biopatina on ornamental fountains: (a). Fountain located in the Royal Alcazar of Seville. (b) Fountain located in the Alhambra and Generalife monument (Granada, Spain).
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Figure 2. Damage caused by biodeterioration on ornamental stone. Fountains located in the English Garden of the Royal Alcazar of Seville, Spain: (a) Discolouration of the stone support; (b) Disintegration and cracking of the support.
Figure 2. Damage caused by biodeterioration on ornamental stone. Fountains located in the English Garden of the Royal Alcazar of Seville, Spain: (a) Discolouration of the stone support; (b) Disintegration and cracking of the support.
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Figure 3. Bioactive compounds and the effects of algicidal bacteria.
Figure 3. Bioactive compounds and the effects of algicidal bacteria.
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Figure 4. Conventional treatment with chlorine tablets (a). Persistence of photosynthetic microorganisms and insufficient efficacy (b). Fountain of Emperator Charles V, (Alhambra, Granada, Spain).
Figure 4. Conventional treatment with chlorine tablets (a). Persistence of photosynthetic microorganisms and insufficient efficacy (b). Fountain of Emperator Charles V, (Alhambra, Granada, Spain).
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Table 1. Freshwater photosynthetic microorganisms, their associated risks to cultural heritage, and reported algicidal bacteria.
Table 1. Freshwater photosynthetic microorganisms, their associated risks to cultural heritage, and reported algicidal bacteria.
Photosynthetic MicroorganismRisks to Cultural HeritageAlgicidal BacteriaReference
Unicellular cyanobacteria
Microcystis sp.1, 2, 4, 5, 6, 7, 8, 9, 11, 12Against Microcystis aeruginosa:
Achromobacter sp.
Acinetobacter sp.
Aeromonas sp.
Aeromonas bestarium
Aeromonas guillouiae
Aeromonas veronii
Agrobacterium vitis
Aquimarina salinaria sp. nov
Aquimarina sp.
Bacillus amyloliquefaciens
Bacillus cereus
Bacillus fusiformis
Bacillus licheniformis
Bacillus methylotrophicus
Bacillus mycoides
Bacillus thuringiensis
Bacillus siamensis
Bacillus sp.
Bacillus subtilis
Bacillus pumilus
Brevibacillus sp.
Brevundimonas diminuta
Chryseobacterium sp.
Cytophaga sp.
Enterobacter sp.
Enterobacter hormaechei
Exiguobacterium sp.
Halobacillus sp.
Leuconostoc mesenteroides
Lysobacter sp.
Morganella morganii
Ochrobactrum sp.
Paebubacillus sp.
Paenibacillus polymyxa
Paenibacillus alvei
Paucibacter aquatile
Pedobacter sp.
Pseudomonas sp.
Pseudomonas aeruginosa
Pseudomonas putida
Pseudomonas stutzeri
Pseudomonas syringae
Raoultella planticola
Raoultella ornithinolytica
Raoultella sp.
Rhizobium sp.
Rhodococcus sp.
Serratia marcescens
Saprospira albida
Stenotrophomonas sp.
Stenotrophomonas acidaminiphila
Streptomyces globisporus
Streptomyces eurocidicus
Streptomyces lushanensis sp.nov.
Streptomyces neyagawaensis
Streptomyces sp.
Xanthobacter autotrophicus
Against M. viridis:
Aeromonas sp.
Bacillus cereus
Bacillus sp.
Chryseobacterium sp.
Exiguobacterium sp.
Pseudomonas sp.
Pseudomonas putida
Stenotrophomonas sp.
Streptomyces lushanensis sp.nov.
Against M. wesenbergii:
Bacillus cereus
Bacillus licheniformis
Enterobacter sp.
Stenotrophomonas sp.
Streptomyces lushanensis sp.nov.
Pseudomonas putida
Against M. flos-aquae:
Aeromonas sp.
Streptomyces lushanensis sp.nov.		[7,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78,79,80,81,82,83,84,85,86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103]
Chroococcus sp.1, 2, 3, 6, 7, 9, 11, 13, 14Aeromonas sp.
Bacillus sp.
Chryseobacterium sp.
Exiguobacterium sp.		[46,50,54,66,104]
Filamentous cyanobacteria
Anabaena sp.1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12Pseudomonas sp.
Serratia marcescens, Streptomyces sp.
Streptomyces eurocidicus
Bacillus amyloliquefaciens
Against Anabaena variabilis:
Aeromonas sp.
Lysobacter sp.
Rhodococcus sp.
Pseudomonas sp.
Against Anabaena cylindrica:
Aeromonas sp.
Lysobacter sp.
Pseudomonas sp.
Pseudomonas putida
Against Anabaena flos-aquae:
Aeromonas sp.
Bacillus cereus
Bacillus thuringiensis
Cytophaga sp.
Pseudomonas putida
Streptomyces globisporus
Streptomyces lushanensis sp.nov.		[39,52,53,58,71,81,87,90,93,95,96,99,105]
Leptolyngbya sp.1, 2, 3, 4, 5, 6, 7, 8, 9, 11Pseudomonas fluorescens[106]
Nostoc sp. 1, 2, 3, 4, 5, 6, 7, 8, 9, 11, 12 Bacillus cereus (against N. punctiforme specie)
Bacillus amyloliquefaciens
Exiguobacterium sp.
Streptomyces lushanensis sp. nov.
Pseudomonas fluorescens		[40,54,58,96,106]
Oscillatoria sp.1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 15Aeromonas sp.
Bacillus sp.
Bacillus cereus
Bacillus flexus
Chryseobacterium sp.
Enterobacter sp.
Exiguobacterium sp.
Flexibacter sp.
Pseudomonas sp.
Pseudomonas fluorescens
Stenotrophomonas sp.
Streptomyces globisporus
Against O. tenuis:
Bacillus cereus
Enterobacter sp.
Enterobacter asburiae
Pseudomonas simiae oli
Streptomyces lushanensis sp. nov.
Against O. planctonica:
Bacillus cereus
Streptomyces lushanensis sp. nov.
Exiguobacterium sp.		[41,46,50,54,66,92,93,97,101,106,107,108,109,110]
Phormidium sp.1, 2, 3, 4, 5, 7, 9Bacillus licheniformis
Myxococcus xanthus (against P. luridum specie)
Streptomyces globisporus
Pseudomonas fluorescens
Aquimarina sp. (against P. persicinum specie)		[61,93,106,111,112]
Pseudanabaena sp.1, 2, 3, 4, 6, 7, 8, 9, 10, 11, 12Streptomyces sp.
Pseudomonas fluorescens
Lysobacter cf. brunescens (against limnetica specie)		[98,106,113]
Unicellular green algae
Chlamydomonas sp.1, 2, 4, 7, 8, 9Aeromonas sp.
Aquimarina sp. (against C. raudensis)
Bacillus cereus (against C. reinhardtii specie)
Exiguobacterium sp.
Chryseobacterium sp.
Stenotrophomonas sp.		[46,50,54,92,114]
Chlorella sp.1, 2, 4, 5, 6, 7, 8, 9, 11, 15Bacillus fusiformis
Bacillus sp.
Against C. vulgaris:
Aeromonas sp.
Aquimarina sp.
Bowmanella denitrificans
Enterobacter sp.
Flammeovirga yaeyamensis
Flavobacterium aquatile
Microbacterium paraoxydans
Pseudomonas oleovorans
Stenotrophomonas sp.
Against C. ellipsoidea:
Bacillus cereus
Pseudomonas putida
Against C. emersonii:
Flavobacterium aquatile
Pseudomonas oleovorans
Against C. autotrophica
Streptomyces sp.
Deinococcus sp.
Against C. pyrenoidosa
Aquimarina sp.		[40,42,51,52,55,92,99,101,115,116,117,118,119,120,121]
Choricystis minor2, 4, 5, 6, 7, 8, 9Microbacterium sp.[122]
Scenedesmus sp.1, 2, 3, 4, 5, 6, 7, 11, 15Against S. quadricauda:
Aeromonas sp.
Enterobacter asburiae
Pseudomonas simiae oli
Against S. obliquus:
Bacillus fusiformis 		[49,51,55,107,123]
Filamentous green algae
Spyrogira gracilis1, 2, 4, 7, 8, 9, 11Bacillus subtilis[124]
Diatoms
Synedra acus subsp. radians (Ulnaria)1, 2, 4, 5, 7, 8, 9, 11
* Although Synedra acus subsp. radians has not been reported in cultural heritage biodeterioration, species of the genus Synedra (currently included in Ulnaria) have been observed forming biofilms on stone surfaces in ornamental fountains and similar water-contact environments.		Brevundimonas bullata
Sphingomonas rhizogenes
Agrobacterium tumefaciens
Methylobacterium adhaesivum
Acinetobacter johnsonii
Bacillus simplex
Bacillus mycoides
Deinococcus aquaticus		[125,126]
* Risks to cultural heritage: (1) Biofilm formation; (2) Exopolysaccharide (EPS) production; (3) Pore penetration and enlargement (endolithic/chasmoendolithic behavior); (4) Mechanical stress (hydration/desiccation cycles); (5) Carbonate precipitation and crust formation; (6) Mineral dissolution by metabolic byproducts; (7) Esthetic alteration (e.g., pigmented patinas/crusts, discoloration); (8) Contaminant trapping and accumulation; (9) Facilitation of secondary colonization (fungi, lichens, mosses, etc.); (10) Toxin production (e.g., anatoxin-a, saxitoxin); (11) Water quality deterioration; (12) Health risk for humans and animals; (13) Recolonization after cleaning; (14) Resistance to biocontrol treatments; (15) Post-treatment recolonization potential.
Table 2. Algicidal bacteria: reported bioactive compounds, classification and references.
Table 2. Algicidal bacteria: reported bioactive compounds, classification and references.
Algicidal BacteriaBioactive CompoundsTypeReference
Aeromonas sp.3-methylindole and 3-benzyl-piperazine-2,5-dioneAmino acid and peptide derivatives[46]
Aeromonas guillouiae4-hydroxyphenethylamineAlkaloid[156]
Aeromonas veroniilumichrome, 9-chlorolumichrome, veronimide, and veronipyrazineFlavin derivatives, nitrogen compounds, and pyrazine[48,157]
Aquimarina sp.l-amino acid oxidase (l-AAO)Enzimes[114]
Bacillus amyloliquefaciensBacilysin; L-lysine (Lys) and L-phenylalanine (Phe)Amino acids[58,59,60]
Bacillus cereusN-phenethylacetamide; Ciclo (L-Pro-L-Val); Ciclo (L-Pro-L-Pro)Derivatives of simple amides and cyclic diketopiperazine[158]
Bacillus siamensistryptoline (C11H12N2)Alkaloids[65]
Bacillus sp.
3-Isopropyl-hexahydropyrrolo [1, 2-a] pyrazine-1,4-dione and Hexahydropyrrolo [1,2-a] pyrazine-1, 4-dione
Indole-3-carboxaldehyde, cyclo(Pro-Phe), and unidentified high-molecular-weight compound(s) (>3 kDa)
Alkaloid and diketopiperazine
alkaloid, DKP, and macromolecular compound (>3 kDa)		[66,159]
Bacillus subtilisBacteriocins, sactibiotics, non-ribosomal polypeptides, and lipopeptides (such as surfactin and fengycin) with unidentified antimicrobial activity against freshwater photosynthetic microorganisms Peptides and lipopeptides[160]
Bacillus thuringiensisPurine derivative identified as C12H15O5N5Derivative of nitrogenous bases[70]
Chryseobacterium sp.cyclo(4-OH-Pro-Leu), cyclo(Pro-Leu)Diketopiperazines[50]
Deinococcus sp.DeinoxanthinTerpenes[121]
Enterobacter hormaecheiProdigiosinAlkaloids [72]
Flammeovirga yaeyamensisAmylases, celulases, xylanasesEnzimes[119]
Leuconostoc mesenteroidesphenyl-lactic acidOrganic acids derived from amino acids[75]
Lysobacter sp.L-tyrosineDerivated from amino acids[113]
Microbacterium paraoxydansAtrazine-desethylNitrogenous organic compound[117]
Raoultella sp.dissolved microbial metabolites and humic acid) and smaller amounts of other substances (protein-like substances and fulvic acid) Undefined microbial metabolites and humic substances[161]
Raoultella ornithinolyticaD-Gluconic acid, Chlorogenic acid, L-Malic acid, 5-Hydroxy-2,4-dioxopentanoate, 2-Methyl-3-oxopropanoic acidLow-molecular-weight organic acids (e.g., gluconic acid, malic acid) and a phenolic compound (chlorogenic acid)[84]
Serratia marcescensProdigiosinAlkaloids[89]
Stenotrophomonas sp.Cyclo-(Gly-Pro), HydroquinoneDiketopiperazine and phenolic compound[92]
Streptomyces sp.Flavonoids and unidentified extracellular metabolitesFlavonoids[162]
Streptomyces eurocidicustryptamine and tryptoline Alkaloids[65]
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Calvo-Bayo, I.; Bolívar-Galiano, F.; Romero-Noguera, J. Algicidal Bacteria: A Sustainable Proposal to Control Microalgae in the Conservation and Restoration of Stone Cultural Heritage. Sustainability 2025, 17, 10610. https://doi.org/10.3390/su172310610

AMA Style

Calvo-Bayo I, Bolívar-Galiano F, Romero-Noguera J. Algicidal Bacteria: A Sustainable Proposal to Control Microalgae in the Conservation and Restoration of Stone Cultural Heritage. Sustainability. 2025; 17(23):10610. https://doi.org/10.3390/su172310610

Chicago/Turabian Style

Calvo-Bayo, Isabel, Fernando Bolívar-Galiano, and Julio Romero-Noguera. 2025. "Algicidal Bacteria: A Sustainable Proposal to Control Microalgae in the Conservation and Restoration of Stone Cultural Heritage" Sustainability 17, no. 23: 10610. https://doi.org/10.3390/su172310610

APA Style

Calvo-Bayo, I., Bolívar-Galiano, F., & Romero-Noguera, J. (2025). Algicidal Bacteria: A Sustainable Proposal to Control Microalgae in the Conservation and Restoration of Stone Cultural Heritage. Sustainability, 17(23), 10610. https://doi.org/10.3390/su172310610

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