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Review

Gray Mold in Blueberry: Current Research on Pathogenesis, Host Resistance, and Control Strategies

by
Lifeng Xiao
1,
Qiuyue Zhao
2,
Jie Deng
1,
Lingyan Cui
1,
Tingting Zhang
1,
Qin Yang
1,* and
Sifeng Zhao
2,*
1
Provincial Famous Teacher Yang Qin Studio/Key Laboratory of Molecular Breeding and Variety Creation of Horticultural Plants for Mountain Features in Guizhou Province, School of Life and Health Science, Kaili University, Kaili 556011, China
2
Key Laboratory of Oasis Agricultural Pest Management and Plant Protection Resources Utilization, Shihezi University, Shihezi 832003, China
*
Authors to whom correspondence should be addressed.
Horticulturae 2025, 11(10), 1241; https://doi.org/10.3390/horticulturae11101241
Submission received: 31 August 2025 / Revised: 9 October 2025 / Accepted: 13 October 2025 / Published: 14 October 2025
(This article belongs to the Section Plant Pathology and Disease Management (PPDM))
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Abstract

Gray mold, caused by Botrytis cinerea, poses a significant fungal threat to postharvest blueberries, leading to substantial economic losses and challenging the sustainable development of the blueberry industry. This highlights the urgent necessity for comprehensive research to develop effective and sustainable management solutions. This review offers a systematic overview of gray mold in blueberries, with a particular emphasis on elucidating the pathological mechanisms employed by B. cinerea, including its infection pathways and virulence factors. It examines the resistance mechanisms in blueberries, which include both preformed and induced physical and biochemical defenses, and synthesizes existing control strategies. These strategies range from conventional fungicides to emerging alternatives such as biological control agents, natural antimicrobials, physical treatments, and integrated pest management (IPM) approaches. Furthermore, the paper explores future research directions by identifying key knowledge gaps and promising areas for innovation. This study aims to bridge the gap between fundamental knowledge and practical application, thereby providing a robust theoretical foundation and actionable guidance for the effective prevention and management of gray mold in blueberry production and storage.

1. Introduction

Blueberry (Vaccinium spp.) is esteemed globally as a valuable economic crop, primarily due to its high concentrations of antioxidants, including anthocyanins, phenolic acids, and flavonoids, which have earned it the moniker “king of berries.” According to FAOSTAT, global blueberry production has reached over 1.7 million metric tons, highlighting the crop’s major agricultural significance. Gray mold, caused predominantly by Botrytis cinerea Pers., is one of the most commonly observed and severe diseases of blueberry worldwide, particularly in the post-harvest stage where it leads to significant economic losses, and is well documented in countries such as Canada, Korea, Chile, and China [1,2]. Given the growing global demand for blueberries, understanding the pathogen’s biology and the plant’s defense responses is essential for developing sustainable control strategies. This review article addresses this problem by synthesizing current knowledge on pathogenic mechanisms, host resistance, available management practices, and potential research perspectives, thereby contributing to both scientific understanding and practical application in blueberry production systems.
Gray mold is among the most critical fungal diseases impacting blueberry after harvest [3]. Gray mold is not only prevalent in blueberries but also affects a variety of other economically significant crops, including grapes, tomatoes, tobacco, cucumbers, and strawberries [4,5,6,7]. B. cinerea is a necrotrophic filamentous fungus with a broad host range, affecting more than 1400 plant species, thus becoming one of the main threats to the agriculture systems [8,9,10]. Its conidia can reach host tissues via air currents, water droplets, or mechanical damage. The fungus remains active in low temperature (0–4 °C) and humid conditions, leading to gray mold incidence levels as high as 60–100% during the post-harvest storage of blueberry [11].
Upon infection with gray mold, blueberry exhibit characteristic symptoms on the fruit, flowers, and leaves. Initially, the fruit develops small, hard, light brown lesions that rapidly expand and become covered with a layer of gray fungal growth. Flowers display symptoms of rot, with severe infections leading to complete wilting. Leaves may form brown to gray lesions, and under conditions of high humidity, gray mycelium and masses of conidia may form on these lesions [12]. Gray mold not only directly causes rot in blueberries but also diminishes their shelf life by accelerating water loss and nutrient degradation. B. cinerea, when cultured on potato dextrose agar (PDA) medium, initially forms white colonies that subsequently transition to gray or brown and produce black sclerotia. The conidiophores are slender and branched, while the conidia are oval to round in shape [13,14].

2. Pathological Mechanism of Gray Mold in Blueberry

B. cinerea, a facultative parasite, compromises host cell integrity through the secretion of cell wall-degrading enzymes, including pectinase and cellulase, as well as toxins such as oxalic acid, while concurrently suppressing plant immune responses [15]. At the cellular level, the metabolic byproducts of this fungus cause damage to blueberry cells. Research suggests that the metabolic phenotypes of B. cinerea demonstrate significant similarity across various crops, indicating its strong adaptability to different hosts. This adaptability may be attributed to its extensive metabolic capabilities, including the ability to utilize multiple sources of carbon, nitrogen, phosphorus, and sulfur [16]. These metabolic characteristics provide a theoretical basis for understanding the biochemistry and metabolic phenotypes of B. cinerea, offering valuable insights for the management of gray mold.
The susceptibility of blueberry to gray mold varies throughout different developmental stages, with research indicating that the flowering and fruiting stages, particularly during peak flowering (full bloom) and mature fruit stages, are most vulnerable to infection by B. cinerea [17]. Geographically, blueberry gray mold is prevalent across all regions where blueberries are cultivated. Notably, in certain areas such as Florida, USA, blueberry planting proximity to strawberry fields exhibit higher incidence levels of gray mold [18]. Furthermore, climatic conditions play a crucial role in the prevalence of blueberry gray mold. Empirical evidence suggests that when humidity persists for more than 6 h and temperatures range from 14 to 25 °C, the likelihood of gray mold infection increases, displaying a significant positive correlation with the incidence of gray mold in stored fruit [17].

3. Resistance Mechanisms of Blueberries to Gray Mold

Understanding the resistance mechanisms of blueberry to gray mold is essential for the sustainable advancement of the blueberry industry. Recent studies have explored these mechanisms from various angles, offering a theoretical foundation for future research and application.
The epidermal structure of blueberry, along with certain secondary metabolites, contributes to their defense against fungal infections. Research on cuticular components, particularly ursolic acid (UA), shows its ability to damage B. cinerea membranes, which is important for understanding cultivar differences (as the concentration and composition of cuticular wax, including UA, can vary significantly among cultivars) and potentially enhancing storage shelf life. The antifungal action of UA involves inducing ROS accumulation and inhibiting ROS scavenging enzymes, increasing oxidative stress in B. cinerea cells and damaging their membranes [19]. The study underscores the importance of cuticular components in cultivar resistance and storage, suggesting that understanding their biochemical interactions with the pathogen can lead to targeted fruit preservation strategies. This research advances our knowledge of plant-pathogen interactions and opens avenues for innovative post-harvest treatments using fruits’ inherent defenses.
From a physiological and biochemical standpoint, the resistance of blueberry to gray mold involves the participation of multiple enzymes and secondary metabolites. As shown in Figure 1, research indicates that methyl jasmonate (MeJA) treatment markedly enhances the activity of antioxidant and defense-related enzymes in blueberry, including superoxide dismutase, catalase, ascorbate peroxidase, chitinase, and β-1,3-glucanase, while concurrently decreasing membrane lipid peroxidation levels. Additionally, MeJA treatment augments the phenylalanine pathway, leading to increased activity of enzymes such as phenylalanine ammonia-lyase, cinnamic acid 4-hydroxylase, and 4-coumaroyl-CoA ligase, which in turn elevates the levels of secondary metabolites such as phenolic compounds and flavonoids [20]. Similarly, treatment with salicylic acid-grafted bamboo hemicellulose (HC-g-SA) has been shown to activate the phenylpropanoid metabolic pathway, enhance the accumulation of lignin and flavonoids, reduce the incidence of gray mold by 40%, and modulate the microbial community on the fruit peel to suppress pathogen abundance [21].
Additionally, melatonin has been shown to enhance post-harvest disease resistance in blueberry by modulating the jasmonic acid signaling pathway and the phenylpropanoid pathway. This modulation results in the upregulation of related gene expression and promotes the accumulation of total phenols, flavonoids, anthocyanins, and lignin [22].
Regarding the regulation of signaling molecules, MeJA is pivotal in inducing resistance in blueberry against gray mold. Research has demonstrated that MeJA treatment can induce 782 differentially expressed metabolites, activate the phenylpropanoid metabolic pathway (leading to the accumulation of rutin and quercetin), and promote the synthesis of unsaturated fatty acids (resulting in increased linolenic acid), thereby providing carbon skeletons and energy [23] (Figure 1). Empirical evidence indicates that MeJA treatment triggers a surge in nitric oxide (NO) and elevates endogenous hydrogen peroxide (H2O2) levels in blueberry. These signaling molecules are instrumental in activating resistance responses in blueberry to combat B. cinerea infection. Moreover, NO and H2O2 exhibit a hierarchical relationship in the MeJA-induced resistance mechanism, with NO functioning upstream of H2O2 [20,24] (Figure 1)). The findings indicate that MeJA may enhance the resistance of blueberries to B. cinerea by modulating antioxidant and defense-related enzymes, as well as the phenylpropanoid pathway, through the activation of signaling molecules.
In tomatoes, the expression of the jasmonic acid transporter SlABCG9 is markedly up-regulated in response to gray mold infection [25]. The transcription factor SlbHLH95 is integral to the regulation of flavonoid metabolism and the enhancement of gray mold resistance in tomato fruits. It achieves this by modulating key enzyme genes, thereby promoting flavonoid accumulation and enhancing resistance to gray mold through the suppression of the expression of SlBG10, a β-1,3-glucanase that breaks down callose [26]. Additionally, the plant autophagy mechanism has been shown to significantly contribute to gray mold resistance. In grapevines, VvATG18a, a target of the brassinosteroid (BR) signaling pathway, enhances resistance to gray mold by modulating the autophagy pathway. Furthermore, the exogenous application of the BR plant hormone 24-epibrassinolide (eBR) has been found to activate the BR signaling pathway, thereby augmenting grape disease resistance [27].
In cucumbers, polygalacturonase-inhibiting protein (PGIP) has been shown to limit the growth and colonization of B. cinerea, thereby bolstering the plant’s immune function. Research indicates that the expression of Cucumis sativus polygalacturonase-inhibiting protein 2 (CsPGIP2) is upregulated during gray mold infection and may interact with en-dopolygalacturonase 3 (BcPG3), a virulence factor, to inhibit the pathogen’s invasive capabilities [28]. Moreover, transgenic Arabidopsis plants expressing CsPGIP2 demonstrated significantly reduced areas of gray mold infection, underscoring the critical role of CsPGIP2 in enhancing plant disease resistance [28].
Collectively, these studies suggest that both the metabolic adaptability of B. cinerea and the immune responses of plants collaboratively influence the pathological mechanisms underlying gray mold disease. This dynamic equilibrium between the pathogen and its host plant dictates the occurrence and progression of gray mold. Thus, thoroughly examining the metabolic traits of B. cinerea and plant immune responses not only helps clarify the disease mechanisms of gray mold but also offers a scientific foundation for creating new disease control methods. Further investigation is required to elucidate the resistance mechanisms of blueberry against gray mold and to develop more effective control strategies. These strategies should integrate biological, physical, and chemical methods to support the sustainable growth of the blueberry industry.

4. Research on the Control of Gray Mold in Blueberry

4.1. Chemical Control

Various fungicides, including benzimidazoles, dimethylimidazoles, triazoles, strobilurins, and succinate dehydrogenase inhibitors (SDHIs) have been employed to manage gray mold in blueberry. Each fungicide operates through distinct mechanisms [29]. For instance, benzimidazole fungicides primarily inhibit the synthesis of microtubule proteins in pathogens, disrupting their mitosis and thereby exerting fungicidal effects [30].
Recent advancements have been made in the development of active ingredients for controlling gray mold. Some new fungicides, such as benzovindiflupyr, demonstrate potent inhibitory activity against gray mold [31]. Salt solutions such as potassium bicarbonate, calcium chelates, and sodium silicate have shown notable inhibitory effects on B. cinerea by altering fungal structure and cellular functions [32]. Other reports of novel compounds with notable activity against B. cinerea include certain derivatives of benzoylurea [33], pyrimidine [34], flavonoid [35], and 1,2,4-triazolo [4,3-c]trifluoromethylpyrimidine [36]; further study of these molecules may lead to the development of new fungicides against gray mold. Most recently, researchers developed a range of aldehyde-thiourea derivatives, and tests showed that some compounds had moderate to excellent antifungal effects against gray mold, surpassing current fungicides [37].
The emergence of resistance to multiple fungicides poses a substantial challenge to the management of blueberry gray mold [30] as the prevalence of fungicide resistance in B. cinerea continues to escalate [29]. For instance, a study conducted in Michigan reported an increasing frequency of resistance to multiple fungicides, with reduced sensitivity frequencies particularly noted for pyraclostrobin and difenoconazole [38]. Similarly, in the Pacific Northwest, B. cinerea has developed resistance to SDHI fungicides, with certain mutations resulting in cross-resistance to various SDHI active ingredients [39]. It underscores the necessity for balanced fungicide use and rotation to manage resistance effectively.
A comprehensive study examined the sensitivities of 249 California and 106 Wash- ington B. cinerea isolates to five fungicides from different classes (boscalid, cyprodinil, fenhexamid, fludioxonil, and pyraclostrobin) [40]. The risk of resistance to QoI fungicides was assessed, indicating a higher resistance risk in California. On detached blueberry fruit, most fungicides failed against resistant isolates of the fungus, but a cyprodinil and fludi-oxonil mixture was effective against all tested resistant phenotypes [40].
Addressing these challenges necessitates the optimization of chemical control strategies and the development of novel pesticides. Peroxyacetic acid (PAA), recognized for its low risk to human health, holds promising potential for post-harvest treatment of blueberries. Empirical studies have demonstrated that immersing blueberries in a PAA solution at a concentration of 85 μL·L−1 effectively reduces fruit rot incidence, while maintaining fruit firmness and the sugar-to-acid ratio, and without compromising fruit quality or sensory attributes [41]. PAA has been extensively utilized in the post-harvest management of diseases in citrus and grapes, where the antifungal activity of chlorine dioxide (ClO2) and PAA under both in vivo and in vitro conditions was similar to the chemical fungicides. At lower concentration, ClO2 was more active than PAA [42]. The underlying mechanism is hypothesized to involve the disruption of pathogen cell membrane integrity. In parallel, natamycin, a naturally occurring antifungal compound that disrupts fungal membrane function through ergosterol binding, has been shown by Saito et al. (2022) to significantly decrease the incidence of blueberry gray mold caused by fungicide-resistant B. cinerea at a concentration of 0.46 g/L, without adverse effects on sensory quality, and also reduced overall fruit decay caused by natural infections [43].

4.2. Physical Control Measures

Physical control measures predominantly encompass the regulation of temperature and humidity, alongside ultraviolet (UV) irradiation. Empirical studies have demonstrated that periodic adjustments in temperature and humidity can effectively suppress the progression of gray mold in tomatoes, which serve as a model organism for such research [44]. Furthermore, the manipulation of gas composition within storage environments, such as through the use of a graduated controlled atmosphere (gradually reaching 5 kPa O2 and 10 kPa CO2 respectively, 0 ± 0.5 °C), can reduce blueberry respiration metabolism, delay aging, and reduce the occurrence of gray mold [45].
The utilization of surface disinfectants is a vital approach for extending the shelf life of blueberries and preserving their quality. A sustained-release antimicrobial protective coating derived from bamboo xylan shows promise in prolonging the storage duration of blueberries. Studies have demonstrated that this coating significantly reduces the incidence of spoilage while maintaining the texture, flavor, and nutritional integrity of the fruit [46]. Furthermore, ultraviolet-C (UV-C) irradiation technology, a non-chemical disinfection method, has been extensively employed for the surface disinfection of blueberries. Nevertheless, the irregular shape of blueberry fruits poses challenges to the disinfection efficacy of conventional UV-C techniques. Research suggests that the design of UV-LED systems can enhance the antimicrobial effectiveness of UV-C technology, thereby improving surface disinfection for irregularly shaped fruits [47]. Additionally, UV-TiO2 photocatalytic technology has exhibited high bactericidal efficiency in the surface disinfection of blueberries. Compared to UV-C alone, UV-TiO2 photocatalysis offers superior performance in reducing surface microbial counts without leaving chemical residues [48]. Research on surface disinfectants for blueberry storage is progressing towards more efficient and eco-friendly solutions. Optimizing these technologies boosts food safety, enhances storage quality, and offers better preservation for blueberries and other perishable fruits.
The distinctive characteristics of nanomaterials present considerable potential for prolonging the shelf life of blueberries and enhancing their resistance to diseases. The utilization of nanomaterials in the storage of blueberries primarily capitalizes on their antibacterial and antioxidant properties. For example, flower-like zinc oxide (ZnO) mediated by carboxylated cellulose nanocrystals exhibits substantial antimicrobial activity even in the absence of light, successfully extending the shelf life of blueberries to over 21 days [49]. Recent studies have developed a starch-based composite film embedded with curcumin-loaded nanocomposites, which significantly enhances the appearance and nutritional value of blueberries while sustaining high activity levels of various antioxidant enzymes [50]. In a similar vein, chitosan nanocapsules infused with clove essential oil, synthesized through ion crosslinking technology, have been shown to effectively prolong the shelf life of blueberries while preserving their firmness and moisture content [51].
However, the environmental toxicity of nanoparticles is concerning due to their potential for persistence, bioaccumulation, and cellular interactions [52]. Future research endeavors should focus on elucidating the mechanisms underlying the application of nanomaterials in blueberry storage to develop more efficient and environmentally sustainable preservation strategies.

4.3. Biological Control Strategies

In recent years, biological control has emerged as a prominent, environmentally sustainable strategy for managing blueberry gray mold.

4.3.1. Antagonistic Microorganisms

Various microorganisms, including Bacillus, Streptomyces, Pseudomonas spp., and Candida oleophila, are known to control gray mold in crops. They achieve biocontrol of postharvest gray mold through nutrient competition, hyphal parasitism, induced systemic resistance, and natural antagonistic compounds [1]. Notably, the Bacillus siamensis YJ15 strain, isolated from blueberries, has been found to produce volatile organic compounds (VOCs) that effectively inhibit B. cinerea and 11 other plant pathogenic fungi. With increased fermentation time, this strain significantly reduces both the incidence and lesion diameter of post-harvest gray mold in blueberry [53], highlighting its substantial potential for biological control as a new and environmentally friendly alternative for postharvest pathogen control.
Additionally, strains of Bacillus velezensis (BA3 and BA4) and Asaia spathodeae (BMEF1), isolated from blueberry plantings, have shown antagonistic effects against B. cinerea and Alternaria alternata in both in vitro and in vivo experiments, indicating their potential as biological control agents for local blueberry production [54]. Moreover, Rouxiella badensis SER3, although originally isolated from strawberry leaves, has exhibited varying levels of antagonistic activity against 20 fungal pathogens affecting berries, including blueberry, and has demonstrated a strong inhibitory effect on the growth of B. cinerea [55]. Furthermore, Burkholderia contaminans 128 was isolated from the roots of blueberry bushes, and its cell-free supernatant was found to effectively inhibit the mycelial growth and conidial germination of B. cinerea, thereby reducing gray mold in blueberries by approximately 90% without compromising fruit quality [56].
Antagonistic yeasts such as Trichosporon sp. and Cryptococcus albidus show strong biocontrol potential against gray and blue molds in apples and pears, effectively inhibiting pathogen growth in low-temperature and controlled environments [57]. Similar studies on the biological control efficacy and potential mechanisms of isolated epiphytic yeasts against post-harvest gray mold in blueberry are needed. A recent study identified three yeast species—Papiliotrema terrestris, Hanseniaspora uvarum, and Rhodosporidium glutinis—as demonstrating significant control efficacy against post-harvest diseases in blueberry, with H. uvarum exhibiting the highest efficacy. This yeast species directly inhibits the pathogen by suppressing spore germination and mycelial growth, as well as producing antifungal VOCs [58].
Regarding control mechanisms, various biological control agents possess distinct characteristics. The fermentation broth and cell-free supernatant of Bacillus tequilensis KXF6501 effectively inhibit the germination of conidia and the mycelial growth of B. cinerea by activating the phenylpropanoid metabolic pathway in blueberry. This activation enhances the activity of related enzymes and stimulates the synthesis of lignin, total phenols, and flavonoids, thereby effectively managing blueberry gray mold [59]. Interestingly, certain amphibian skin bacteria have shown strong antifungal activity against plant pathogens, including B. cinerea. These bacteria mitigate the activity of B. cinerea by modulating plant transcriptional responses, inhibiting fungal growth under in vitro conditions, and significantly decreasing the incidence of gray mold when applied to blueberry [60]. As another example, metabolic products generated by the entomopathogenic soil fungus Metarhizium anisopliae can compromise the cell membrane integrity of B. cinerea, resulting in the leakage of intracellular substances and subsequently inhibiting the pathogen’s growth [61].
RNA interference-based biofungicides are regarded as a promising next-generation strategy for the control of gray mold. By targeting critical fungal genes with double-stranded RNA (dsRNA), the pathogenicity of gray mold fungi can be effectively suppressed. Spray-induced gene silencing (SIGS) technology has shown potential in mitigating gray mold infections on strawberry and tomato fruits [62]. These studies provide both theoretical foundations and practical guidance for the development of more efficient control measures in the future.

4.3.2. Plant-Derived Antimicrobial Substances

Beyond the utilization of microorganisms, plant extracts have demonstrated some potential in the biological control of blueberry gray mold. Research indicates that diterpenoid compounds found in sunflower head extracts possess notable antifungal properties against B. cinerea, with specific compounds showing substantial inhibitory effects. The ethyl acetate extract from sunflower heads protected 42.9% of tested blueberry fruits from B. cinerea infection at a concentration of 1.6 mg/mL, indicating its potential as a biological control for mitigating postharvest diseases in fruits [63].
One study explored the inhibitory effects and mechanisms of Artemisia annua (sweet wormwood) essential oil (AAEO) against three fungi, including the pathogen responsible for blueberry gray mold. AAEO demonstrated significant antifungal activity against these fungi through non-contact exposure, with mechanisms involving the alteration of cell wall and membrane integrity, reduction in biofilm formation, and decrease in cell viability and respiratory activity. Fumigation with AAEO during blueberry storage markedly reduced both blueberry rot incidence and pathogen infection [64]. Research suggests that essential oil volatiles not only effectively manage diseases but also positively influence the storage quality of fruit by reducing fruit weight loss and maintaining ascorbic acid and carotenoid content [65]. Thus, under certain conditions, the application of essential oils can effectively control fruit diseases, offering a potential environmentally friendly solution for the management of blueberry gray mold. Future research should further explore synergistic effects among different biological control techniques to enhance their practical application efficiency in fruit disease management.
Overall, substantial advancements have been achieved in the biological control of blueberry gray mold, with various microorganisms and plant extracts exhibiting promising efficacy and potential for disease management. Nevertheless, the effectiveness of biological control is subject to a multitude of factors. Environmental conditions play a crucial role in influencing the activity and efficacy of biological control agents. Research indicates that certain biological control agents experience diminished activity under conditions of elevated temperature and humidity, resulting in reduced control efficacy [66]. Moreover, the mechanisms of action of biological control agents are relatively complex, and their interactions with pathogens may be disrupted by other microorganisms, thereby impacting their control efficacy. Furthermore, variations in pathogen population structures and ecological environments across different regions may result in inconsistent efficacy of biological control agents, thereby limiting their widespread application and promotion [67]. Consequently, further in-depth research is necessary to facilitate the extensive application of biological control technology for managing blueberry gray mold disease.

5. Integrated Control and Sustainable Management

It is imperative to prioritize cultural control measures in blueberry cultivation. These include ensuring optimal air flow and light conditions in blueberry plantings, promptly removing diseased plant debris, and minimizing the proliferation and dissemination of pathogens. Furthermore, appropriate pruning of blueberry plants and regulation of plant density are crucial for reducing humidity and creating an environment that is unfavorable for the development of gray mold [30].
The integration of chemical and biological control strategies is also widely adopted. For instance, during the blueberry flowering period, biological control agents such as B. tequilensis can be employed for preventive measures, while in the early stages of disease manifestation, the application of low-dose chemical fungicides in conjunction with biological agents can be effective. Research demonstrates that this integrated approach not only effectively manages gray mold but also reduces the reliance on chemical fungicides and mitigates the risk of resistance development [59]. Furthermore, natural compounds such as chitosan, which dissolve in acidic solutions and demonstrate specific control effects against gray mold, can be integrated with other management strategies to improve overall control efficacy [68].
Sustainable management approaches prioritize long-term, environmentally friendly control principles. On one hand, the selection and breeding of blueberry varieties with high resistance to gray mold can fundamentally reduce the incidence of the disease. This can be achieved by studying wild blueberry resources to identify disease-resistance genes and incorporating these genes into cultivated varieties through hybrid breeding techniques [69]. On the other hand, ecological regulation methods can be utilized to maintain the ecological balance of orchards. For example, exploiting antagonistic relationships between microorganisms by introducing beneficial microbial communities, such as certain Bacillus species that produce antimicrobial substances, can be an effective strategy. Furthermore, enhancing the management of orchard soil to preserve its fertility and structure can bolster the intrinsic resistance of blueberry plants, thereby promoting the sustainable management of blueberry gray mold [70].
Integrated Pest Management (IPM) seeks to amalgamate multiple effective control strategies into an efficient and practical management framework. During various stages of blueberry growth, control measures can be strategically implemented based on the risk and characteristics of disease manifestation. For instance, post-harvest, the application of biological control agents, such as Simplicillium lamellicola BCP, can be integrated with physical control methods, including low-temperature storage and ultraviolet irradiation, to significantly enhance the efficacy of managing blueberry gray mold. Studies have demonstrated that the wettable powder formulation BCP-WP10, derived from S. lamellicola BCP, achieves control efficacies of 64.7% and 82.6% against gray mold in tomatoes and ginseng, respectively, when diluted at 500-fold and 250-fold concentrations. When combined with low-temperature storage and UV irradiation, the control efficacy is further enhanced [71].
There are few chemical options for controlling gray mold after harvest. The development and application of forecast systems is essential. A Chilean model for gray mold control on blueberries was inaccurate in South Africa, prompting the development of a local model using hourly weather data. Tested in 2020 on two Western Cape farms, this model required similar or fewer sprays than a standard schedule and resulted in similar or lower gray mold incidence. The study demonstrated that using the decision support system (DSS) provided effective gray mold control with equal or reduced fungicide use [72]. Application of hyperspectral imaging (HSI) was investigated for early and non-destructive detection of B. cinerea infection [73]. The integration of HSI and chemometric analysis exhibited significant potential for rapid and non-destructive evaluation of fruits infected with fungi during storage.
Based on the aforementioned research, a decision-making framework for preventing gray mold during blueberry storage is proposed in Figure 2.

6. Challenges and Future Perspectives

Substantial research advancements have been achieved in pathogen identification, pesticide development, biological control, and the utilization of natural plant extracts for managing blueberry gray mold. Nevertheless, further research is imperative to elucidate pathogenic mechanisms and to develop more efficient and environmentally sustainable control measures to support the healthy progression of the blueberry industry.
Future research directions in the realm of blueberry gray mold control technology will predominantly concentrate on the following areas: firstly, there is a need for further exploration of novel biological control resources. Nature harbors a vast array of microorganisms and natural compounds with antimicrobial properties. It is feasible to discover more effective and safer biological control agents through systematic screening and identification processes. For instance, investigating the rhizosphere microbial community associated with blueberry may uncover microorganisms that form symbiotic relationships with the plant and possess the capability to inhibit B. cinerea. Such discoveries could lead to the development of novel biological control products [54].
Concurrently, it is imperative to enhance research on the pathogenic and resistance mechanisms of this pathogen. Utilizing advanced technologies such as genomics, transcriptomics, and proteomics allows for a comprehensive understanding of the pathogenic processes and molecular mechanisms that contribute to resistance development in B. cinerea. This knowledge provides a theoretical basis for devising targeted control strategies. For example, analyzing the gene expression changes in B. cinerea under varying environmental conditions can help identify key pathogenic and resistance genes, offering potential targets for the development of new fungicides or biological control methods [74]. Moreover, advancing precise and efficient detection technologies represents a critical future direction. The development of innovative detection methods leveraging nanotechnology and biosensors is essential for achieving early, rapid, and accurate identification of blueberry gray mold, thereby enabling timely implementation of control measures and enhancing control efficacy [75]. Additionally, the optimization and integration of diverse control technologies to establish a comprehensive, efficient, and sustainable integrated control system is a pivotal focus of future research in the management of blueberry gray mold.
Future research should prioritize understanding the synergistic mechanisms of multiple technologies to achieve effective control of post-harvest diseases in blueberry and support sustainable industrial development.

Author Contributions

Conceptualization, L.X. and Q.Z.; methodology, S.Z. and L.X.; validation, L.C. and J.D.; investigation, T.Z.; writing—original draft preparation, L.X.; writing—review and editing, Q.Y. and S.Z. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Specialized Fund for the Doctoral of Kaili University (grant No. BS20240216), Provincial famous teacher Yang Qin studio (MSGZS SJ-2024002), the Key Laboratory of the Department of Education of Guizhou Province (No. Qianjiaoji [2022] 053), and Guizhou Key Laboratory of Molecular Breeding for Characteristic Horticultural Crops [2025] 027, the Key Discipline Construction Project for Horticulture of Guizhou Province of Kaili University (grant number ZDXK [2014] 28 of Guizhou Academic Degrees Office) and the First-Class Discipline Construction Project for Horticulture of Kaili University (grant number YLXK2021002).

Data Availability Statement

No new data were created or analyzed in this study. Data sharing is not applicable to this article.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
PDApotato dextrose agar
PALphenylalanine ammonia-lyase
C4Hcinnamic acid 4-hydroxylase
MeJAmethyl jasmonate
HC-g-SAsalicylic acid-grafted bamboo hemicellulose
NOnitric oxide
H2O2hydrogen peroxide
PGIPpolygalacturonase-inhibiting protein
SDHsuccinate dehydrogenase
SDHIsuccinate dehydrogenase inhibitors
PAAPeroxyacetic acid
UVultraviolet
VOCsvolatile organic compounds
dsRNAdouble-stranded RNA
SIGSspray-induced gene silencing
AAEOArtemisia annua essential oil
SODsuperoxide dismutase
CATcatalase
MDAmalondialdehyde
NOSnitric oxide synthase

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Horticulturae 11 01241 g001
Figure 1. Methyl jasmonate (MeJA)–mediated resistance to Botrytis cinerea in blueberry.
Figure 1. Methyl jasmonate (MeJA)–mediated resistance to Botrytis cinerea in blueberry.
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Figure 2. An Evidence-Based Decision Guide for Blueberry Storage to Prevent Gray Mold.
Figure 2. An Evidence-Based Decision Guide for Blueberry Storage to Prevent Gray Mold.
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MDPI and ACS Style

Xiao, L.; Zhao, Q.; Deng, J.; Cui, L.; Zhang, T.; Yang, Q.; Zhao, S. Gray Mold in Blueberry: Current Research on Pathogenesis, Host Resistance, and Control Strategies. Horticulturae 2025, 11, 1241. https://doi.org/10.3390/horticulturae11101241

AMA Style

Xiao L, Zhao Q, Deng J, Cui L, Zhang T, Yang Q, Zhao S. Gray Mold in Blueberry: Current Research on Pathogenesis, Host Resistance, and Control Strategies. Horticulturae. 2025; 11(10):1241. https://doi.org/10.3390/horticulturae11101241

Chicago/Turabian Style

Xiao, Lifeng, Qiuyue Zhao, Jie Deng, Lingyan Cui, Tingting Zhang, Qin Yang, and Sifeng Zhao. 2025. "Gray Mold in Blueberry: Current Research on Pathogenesis, Host Resistance, and Control Strategies" Horticulturae 11, no. 10: 1241. https://doi.org/10.3390/horticulturae11101241

APA Style

Xiao, L., Zhao, Q., Deng, J., Cui, L., Zhang, T., Yang, Q., & Zhao, S. (2025). Gray Mold in Blueberry: Current Research on Pathogenesis, Host Resistance, and Control Strategies. Horticulturae, 11(10), 1241. https://doi.org/10.3390/horticulturae11101241

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