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Review

Biostimulants as a Tool for Mitigating Water Deficit Stress in Strawberry Cultivation

by
Júlia Letícia Cassel
1,
Laura Valentina Caus Maldaner
2,
Mateus Possebon Bortoluzzi
1,2,
Luciane Maria Colla
1,
Francisco Wilson Reichert Junior
1,2,
Pedro Palencia
3,* and
José Luís Trevizan Chiomento
1,2
1
Graduate Program in Agronomy, University of Passo Fundo, Passo Fundo 99010-080, RS, Brazil
2
Undergraduate Program in Agronomy, University of Passo Fundo, Passo Fundo 99010-080, RS, Brazil
3
Department of Organisms and System Biology, Polytechnic School of Mieres, University of Oviedo, 33600 Mieres, Asturias, Spain
*
Author to whom correspondence should be addressed.
Agronomy 2025, 15(11), 2643; https://doi.org/10.3390/agronomy15112643
Submission received: 24 October 2025 / Revised: 13 November 2025 / Accepted: 17 November 2025 / Published: 18 November 2025
(This article belongs to the Section Horticultural and Floricultural Crops)
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Abstract

This bibliometric review analyzed research published between 2020 and 2025 addressing water stress in strawberry plants and evidenced the use of biostimulants as a promising tool in mitigating this stress. Water requirement of strawberry plants varies according to the agroecosystem of cultivation and genotype used to establish the crop. Strawberry plants develop large leaves with a high water content and stomata, which results in high transpiration rates. Under water deficit, the photosynthetic capacity of the plant is reduced and the water content in the leaves is lower. Additionally, molecules such as proline, catalase, and peroxidase are produced, indicating enzymatic oxidative stress. Conversely, the fruit quality is positively influenced when the plant suffers water restrictions (up to 75% of the pot/field capacity). The use of biostimulants represents a potential biotool to mitigate water deficit in strawberry plants, such as the application of organic acids, plant extracts, seaweed, bacteria, and fungi. The use of these products in situations of water deficit or aiming at a reduction in water consumption is still a topic of research gaining attention. Therefore, the application of biostimulants combined with irrigation management with lower water consumption corroborates the search for more productive and sustainable agri-food systems.

1. Introduction

The demand for healthy food is growing, and therefore obtaining products with higher nutritional value is a way to improve the health of consumers, but it still represents a challenge for farmers. In addition, the spotlight of consumers is increasingly focused on agri-food systems, which must be environmentally friendly, socially just, technologically adequate, economically viable, and culturally accepted [1].
Strawberries (Fragaria × ananassa Duch.) are key horticultural crops that contribute to agrifood systems. They produce some fruit with high added value that is appreciated for its color, aroma, and flavor. Strawberries are rich in vitamin C and biomolecules with antioxidant activity, such as pelargonidin-3-O-glucoside [2,3]. These metabolites strengthen the immune system, participate in tissue formation, improve iron absorption and help control infections by stimulating the migration of neutrophils to the site of infection to increase phagocytosis and the generation of oxidants, thereby destroying the infectious agent [4]. One of the strategies to enhance the yield and quality of strawberries is the adoption of soilless cultivation systems, in greenhouses, where the incidence of pests and diseases can be reduced and, consequently, the use of agrochemicals [5]. The establishment of strawberry in substrate and greenhouse, in addition to improving the ergonomics of producers, allows for more efficient monitoring and management of irrigation, fertilization, temperature, and plant health. However, a remaining obstacle regarding the management of strawberry plants in this cropping system is related to the sensitivity of this crop to abiotic stress, especially water stress [6], whether in deficit or excess. Additionally, the genotype has a crucial role in the biochemical response to water stress, since transpiration is a physiological feature that is dependent on genotype-environment-cultivation cluster [2,3]. Mild water deficit can improve fruit quality and reduce water waste, enhancing agroecosystem sustainability [3].
Even in greenhouse cultivation, where the microclimate can be more efficiently controlled by producers, water stress can compromise the development of strawberry plants [7]. This abiotic stress can affect the plant’s morphophysiology through reduced photosynthetic capacity, lower chlorophyll production, reduced vegetative growth, altered root development, and the production of lower-quality fruit [8]. One way to mitigate the negative effects of water stress on strawberries is to use biostimulants when cultivating this horticultural crop.
In addition to attenuating the effects of abiotic stresses in several crops, the addition of biostimulants in the growth medium, or directly in the plants, is often related to increased yield and fruit quality of strawberry [9]. However, bioengineering regarding the use of biostimulants as mitigators of the effects of water deficit on strawberry still represents an area requiring further investigation [10]. The lack of information focuses on the types and doses of biostimulants with potential use, at the time of application, in the interaction with cultivars and in the description of their molecular mechanisms and biochemical models [11].
Therefore, through a bibliometric review of the literature, we aim to compile information on the use of biostimulants in strawberry plants, focusing on their effect on the mitigation of water stress in this horticultural crop.

2. Bibliometric Review

To collect information on mitigating strawberry water stress through the use of biostimulants, scientific articles were searched in the Scopus database (https://www.scopus.com/home.uri). For this selection, no filters were applied regarding language or document access. The main findings on this topic of research, published in scientific journals in this field, were considered.
As keywords (language: English), we initially used the term set “fragaria” OR “strawberry” AND “water stress” AND “biostimulant”, which yielded only four documents detected (Figure 1). Then, to expand the search, the dataset was reduced to “fragaria” OR “strawberry” AND “biostimulant”, where 39 scientific articles were found; and “fragaria” OR “strawberry” AND “water stress”, where 51 scientific articles were found (Figure 1).
Of the 90 scientific articles found, 48 were related to water deficit and 37 to the application of biostimulants in strawberry cultivation. During the reading, the article selection criteria were: (i) relationship among the effects of water management on the morphophysiology and yield of strawberry plants; (ii) direct application of biostimulants in this horticultural crop. The five articles that did not meet these criteria were excluded during the selection process. The search for these articles took place from 25 March to 5 April 2025.
All of the articles referenced in the research were published within the last five years (2020–2025), indicating the interest of scientists in deepening this theme with a view to market application. Since 2023 onwards, there was an increase in research related to the application of biostimulants (Figure 2), which may be linked to both plant resilience in the face of abiotic stresses and increased yield and fruit quality. In a climate change scenario involving changes in air and soil humidity and temperature due to excess or insufficient rainfall, these studies become increasingly important. In addition to their effects on parameters such as productivity, the use of biostimulants can impact the biocontrol of pathogens and pests, which can also be exacerbated by climate change.
With regard to the density of publications by country (Figure 3), studies on the utilization of biostimulants in strawberry plants (Figure 3A) are predominantly concentrated in Mexico (14 documents), followed by Italy (10 documents), Greece (9 documents), Australia, South Korea, and Turkey (7 documents), Portugal and Spain (5 documents), and India, the United States of America, and the United Kingdom (4 documents). In the context of the global challenges posed by climate change, water scarcity, and desertification, Mexico has emerged as a notable exemplar in its adoption of advanced strawberry-production technologies and the implementation of alternative systems. These initiatives aim to enhance water efficiency and maintain productivity while addressing water scarcity issues in rural regions [12]. In Italy, the emphasis on organic and sustainable agriculture, in combination with the technification of research aimed at achieving the United Nations Sustainable Development Goals [13], has placed the country at the forefront of research on the application of biostimulants in strawberry plants.
With regard to the research on water stress in strawberry plants (Figure 3B), the majority of studies are concentrated in Turkey (33 documents), followed by China, and the United States of America (the first and second largest producers of strawberries in the world, respectively) (15 documents), Spain (10 documents), Mexico (7 documents), and Iran, Italy, Poland, and South Korea (6 documents). At this juncture, Turkey is worthy of particular notice given the number of studies on water deficit in strawberry plants given that it is the third largest producer of this fruit on a global scale (largest producer in Europe), in addition to facing real problems of water deficit in the producing regions [14]. The country is experiencing an increase in the technification of agricultural production, as evidenced by the incorporation of protected crops and efficient irrigation systems.
After selecting the articles, keyword clusters with at least four occurrences were generated using the VOSviewer software, version 1.6.19 (Figure 4). For the dataset of the research “fragaria” OR “strawberry” AND “biostimulant” (Figure 4A) there were associations between biostimulants and fruit quality, productivity and plant growth, in addition to the association of the species with antioxidant activity and the relationship of abiotic stresses with chitosan. For the set of words “fragaria” OR “strawberry” AND “water stress” (Figure 4B) three clusters were evidenced: (i) association of water stress with controlled studies and photosynthesis; (ii) the association of Fragaria with chlorophyll and botany; (iii) the association of strawberry with irrigation, productivity, growth, and water stress.

3. Strawberry Cultivation Associated with Water Stress

The cultivated strawberry plant, botanically called Fragaria × ananassa Duch., originated from the hybridization between F. virginiana Mill. and F. chiloensis (L.) Mill., in the 1700s [15]. In the realm of small fruits, strawberry plants have emerged as the predominant cultivated species. The aggregate fruit, formed by fruits joined by the floral receptacle, is highly appreciated by consumers due to its color, flavor, aroma, and health-promoting properties [16,17].
The growth and development of strawberry is guided by biogeographic, edaphoclimatic, and ecophysiological factors [15]. Consequently, in cropping systems where environmental factors cannot be effectively controlled, the crop may be exposed to the occurrence of abiotic stresses. The abiotic factors described as influential in strawberry are climate (temperature, humidity), soil (structure, pH, and fertility), water, and sunlight. In addition, these factors are related to each other, with the water deficit being intensified by the increase in temperature, which causes greater evaporation and reduced rainfall [18]. In this sense, environmental conditions lead to many challenges in strawberry production due to shallow root systems and intensive leaf transpiration [19]. In instances where soil and climate requirements are not met, a decline in yield and fruit quality can be observed [16].
Among the abiotic stresses that affect strawberry, drought conditions exert adverse impacts on the physiological and biochemical responses of this crop, affecting its vegetative components, as well as fruit production and quality [20]. Strawberries are extremely sensitive to water deficit. The plant has a superficial root system, in addition to originating large and amphistomatic leaves, with more than 200 stomata mm−2, a higher number in relation to the stomata in the fruits (up to 6 stomata mm−2), and there may be a reduction in the number of stomata in the leaves of plants under water deficit [21,22,23,24,25,26]. In addition, in some phenological stages, such as fruiting, plants have an increased water requirement [27], which makes it even more sensitive to water deficit. In contrast, water scarcity is a global ecological problem, highlighting agriculture as the sector responsible for the largest water consumption [18].
Therefore, the water requirement of strawberry is strongly dependent on climatic conditions, substrate, genotype, and plant phenology [26,28]. Consequently, the data concerning the water requirement of the crop are contrasting in the studies developed, ranging from an average daily intake in a subtropical climate, for the ‘Albion’ cultivar, during the full fruit harvest of 259.6 to 267.7 mL plant day−1 [29]. Furthermore, this volume of water can vary according to the genotype, which plays a crucial role in the biochemical response to water stress, since the transpiration is a physiological feature, dependent on genotype-environment-cultivation cluster [2,3]. In Mediterranean-type climate conditions, characterized by dry and hot summers and moderately cold winters, the demand for water can range from 270 to 420 mm per crop cycle, depending on the water requirement of the cultivar [30]. In the study conducted on the crop coefficient (kc) of the strawberry, it is evident that the average evapotranspiration was 3.8 mm day−1. Furthermore, the total water consumption during the entire growing season was recorded as 873.4 mm for the cultivar ‘Rainha Elisa’ under semiarid climate conditions [31].
The morphophysiological effects of strawberry under water stress (Figure 5, Table 1) include for water deficit: chlorophyll degradation, stomatal closure, changes in the density and size of stomata, suppression of photosynthesis, optimization of the use of photoassimilates, imbalance of reactive oxygen species, reduction in shoot growth, maintenance of root development, reduction in stem diameter, reduction in relative leaf water content, and lower fruit production [8,16,32,33]. In addition, increases in the levels of proline and anthocyanins, which represent physiological mechanisms capable of protecting plants against the effects of stress, are evidenced [34]. On the other hand, the effects of excess water appear in a less evident way in the literature, with the production of more acidic fruits being described [35]. Here, based on the literature consulted (Figure 5, Table 1), the irrigation intensities in strawberry plants were classified, based on pot/field capacity, as: (1) excessive (125%); (2) normal (100%); (3) deficits—mild (75%), moderate (50%), and intense (25%). The experiments that supported this classification (Figure 5) were carried out with cultivation in soil, except for the study by Jiang et al. 2024 [25], in which the strawberry plants were grown in substrate.
From a physiological point of view, the daily transpiration rate of the strawberry plant decreases during water deficit treatment, even though the biomass of the plants increases continuously (initially quickly, then more slowly). In the irrigation dynamics imposing mild, moderate, and intense water deficits, the synthesis of 3-indolebutyric acid (IBA), salicylic acid (SA), abscisic acid (ABA), indole-3-carboxylic acid (ICA), gibberellin A1 (GA1), and jasmonic acid (JA) was increased in moderate drought treatment. The synthesis of SA, 1-aminocyclopropanecarboxylic acid (ACC), and indole-3-acetic acid (IAA) were downregulated in response to severe drought. In rehydration after a severe water deficit, although IBA was downregulated, ACC, IAA, ICA, and isopentenyl adenosine (IPA) were upregulated [25]. This shows an endogenous hormonal response in the plant to severe stresses and, therefore, a more sensitive production of growth and development hormones after rehydration, aiming at the restoration and maintenance of physiology.
From the genetic point of view, under water deficit treatments, all differentially expressed genes were related to binding mechanisms, catalytic activity, metabolic process, and cellular process. The differentially expressed genes for mild deficit (70% of the water content of pot capacity) were associated with mechanisms responsible for pentose and glucose interconversions and sulfur metabolism. These are related to the production of sweeter fruits in plants with mild water scarcity. For moderate deficit (50% of pot capacity), the mechanisms were expressed in the biosynthesis of secondary metabolites and nitrogen metabolism. For severe deficit (20% of pot capacity) the mechanisms were expressed in the pathways of galactose metabolism and fatty acid elongation. When there was a rehydration of the plants that suffered from water deficit, there was a strong influence of the production of ribosomes, biogenesis of ribosomes in eukaryotes, and other pathways. The modulation of the genetic mechanism of the plant under stress can form encodes responsible for producing antioxidant molecules, amino acids, proteins, and other compounds, in a biochemical and physiological defense mechanism [25]. These changes cause direct effects on the morphophysiology of the plant under stress. However, these relationships are not yet clearly described in the literature.
Also, changes in fruit quality are observed under water stress (Table 1). When the plants were subjected to irrigation levels, qualitative variations in the fruit were verified in relation to the contents of sugar, organic acids, and phenolic acids, as well as changes in the levels of ellagic acid, caffeic acid, chlorogenic acid, and catechin. Many of these compounds are beneficial to human health, as they have antioxidant effects acting as an exogenous source of antioxidants to the diet. In this way, it is possible to optimize the appropriate levels of stress that seek to increase these compounds in fruits, without losses in productivity and fruit size. These innovative agricultural strategies and sustainable irrigation management are key to improving fruit quality [16]. In consideration of the challenges posed by diminishing water availability, coupled with the imperative for effective irrigation management, the development of environmental control techniques [33] assumes paramount importance.
In view of this discussion, improving drought tolerance and the underlying mechanisms is crucial to meet the needs of strawberry production with stable yield and high quality [19]. Applications of sodium selenate [39], abscisic acid [16], and 5-aminolevulinic acid [19] have already been tested for the mitigation of the effects of water stress on strawberry plants. Also, considering the growing use of biostimulants to attenuate abiotic stresses, this can be a sustainable and ecological tool to minimize the negative effects of water stress in strawberry.
Table 1. Morphophysiological and quality characteristics of strawberry fruit under water regimes described in the literature (2020–2025).
Table 1. Morphophysiological and quality characteristics of strawberry fruit under water regimes described in the literature (2020–2025).
Parts of a PlantEffects of Water Regimes on Strawberry MorphophysiologyReferences
Root
  • Shallow roots, highly sensitive to water stress.
[25,40,41]
  • Root characteristics may vary among genotypes subjected to water deficit.
[42]
  • Water stress induced by polyethylene glycol reduces fresh and dry root mass.
[43]
  • Changes in nutrient solutions modify the water retention of the substrate, altering the water potential, and interfering with water absorption.
[44]
  • Associated water and thermal stresses caused senescence of old leaves and translocation of nutrients, stimulating the growth of new roots and leaves.
[45]
  • Partial drying of the root zone reduces fruit yield.
[40]
Leaf
  • Water deficit (0.6%) simulated by polyethylene glycol increased the number of leaves and branches;
  • Water deficit (0.9%) increased levels of proline (2.43 mg g−1) and catalase (19.59 AU min g−1).
[46,47]
  • Water deficit simulation with polyethylene glycol reduced the fresh and dry shoot mass, membrane stability, relative water content, and pigment content;
  • With water deficit, there was an increase in the activity of antioxidant enzymes, leaf proline content, soluble sugars, malondialdehyde, and hydrogen peroxide content.
[43]
  • Water stress reduced plant fresh and dry weights, leaf area, chlorophyll content, stomatal conductance, leaf and plant water content, and N, P, K, Ca, and Mg mineral contents in leaves;
  • Reduction of CO2 assimilation and interference in the accumulation of dry matter of the plant;
  • Dehydration of plant caused an increase in membrane permeability, leaf temperature, L-proline, and malondialdehyde contents.
[36,48,49]
  • Water deficit reduces photosynthetic activity and chlorophyll levels.
[23,50]
  • Positive correlation between water regime with plant height (r = 0.80) and leaf area (r = 0.77).
[34]
  • Water deficit (50%) reduced net photosynthesis, stomatal conductance, and transpiration rate;
  • Stomatal closure was the main predictor of decreased photosynthesis.
[38]
  • Deficient irrigation systems can decrease the contents of total chlorophyll, chlorophyll a, chlorophyll b, carotenoids, leaf water potential, and stomatal conductance.
[20,28,51]
  • Water stress increased the levels of superoxide dismutase, catalase, and peroxidase in the shoot.
[52]
Flower
  • The use of biostimulants ensured the maintenance of flower production in plants under water stress.
[53]
  • Water deficit can reduce the number of flowers.
[36]
  • Exogenous proline supplementation in plants under water stress maintained the number of flowers and fruits.
[54]
Fruit
  • Strawberries, in general, had very thin epidermis and high water content.
[55,56]
  • Water deficit reduced fruit length, fruit diameter, and fruit weight.
[34]
  • Reduced water conditions increased soluble solids and citric acid and total phenolic contents of fruits.
[18]
  • Deficient irrigation levels positively influenced total acidity, ellagic acid, caffeic acid, chlorogenic acid, total soluble solids, fructose, glucose, sucrose, and catechin.
  • Water deficit increases sugar levels and accumulation of phenolic compounds.
[16,36]
  • Sustained deficit irrigation can reduce strawberry firmness;
  • Partial drying of the root system (mild water deficit followed by rehydration) increased the anthocyanin indexes, color, and carbohydrate content of the fruit, suggesting an improvement in certain physicochemical characteristics.
[40]
  • Irrigation with water deficit (maintaining up to 75% of the field capacity) increased the yield and the contents of vitamin C, soluble sugar, sugar-acidity ratio, and soluble protein of the fruits.
[37]
  • Fertilization systems with water deficit decreased the concentrations of N, P, and K in the fruits;
  • Additional fertilizers can alleviate the negative impact of water stress on growth, development, and fruit quality.
[20,49,57]
  • Fruit yield ranged from 43.91 to 86.70 g per plant between irrigations from 50 to 100% of the evaporation of the Class A tank;
  • Foliar applications of biostimulants, such as sodium selenate, attenuate the stress of water deficit in terms of fruit yield.
[39]
  • Humidification treatment systems based on vapor pressure deficit increased yield.
[58]
  • Yield
[38]
  • Irrigation systems within the standard or in excess (100 and 125%) increased fruit acidity.
[35]
Stolon
  • Under water stress, the number of stolons per plant increased.
[32]
  • When the stolons are transplanted, given their cut from the mother plant, they need adequate water supply (meeting 100% of the crop’s water demand), given the loss of water through continuous transpiration and incision.
[59]
Hormonal dynamics
  • In water deficit, the plants produced salicylic acid, abscisic acid, 1-aminocyclopropanecarboxylic acid, and indole-3-acetic acid to maintain their normal growth.
[25]
Gene regulation
  • Genes involved in mineral uptake and flavonoid biosynthesis are among the genes inactivated in situations of severe water deficit and can be transcriptionally reversible in case of plant rehydration.
[48]
  • In moderate drought (50% of pot capacity), plant growth and development were improved through the regulation of sugar, N, and S metabolism;
  • In excessive drought (20% of pot capacity), drought resistance was achieved mainly through changes in the synthesis and metabolism of amino acids and fatty acids;
  • Normal plant development and growth were restored during the rehydration treatment after drought, through changes in the nucleotide metabolic pathway.
[25]
  • Temperature coordination genes (Hsp70 and Hsp90), as well as small heat shock proteins (sHps), play a vital role in the tolerance of water stress associated with thermal stress.
[60]

4. Biostimulants and Perspectives in the Attenuation of Water Deficit in Strawberry

4.1. Overview of the Use of Biostimulants

One strategy for developing and maintaining sustainable agrifood systems is to use biostimulants on cultivated plants such as strawberries. In addition to environmentally friendly effects, such as the reduction in greenhouse gas emissions [61], this establishes a balance between yield and fruit quality [62,63]. Plant biostimulation involves applying physical, chemical and/or biological stimuli to achieve a positive response in terms of growth, development and product quality [64].
These bioinputs can generally be classified as either non-microbial biostimulants (humic substances, complex organic materials, beneficial chemical elements, inorganic salts, seaweed extracts, chitin and chitosan derivatives, peptides, amino acids, and other substances containing N, phytoregulators, and phytohormones) or microbial biostimulants (biological products containing living microorganisms) [65] (Figure 6).
The use of biostimulants in strawberry plants aims to improve fruit production and quality, as well as tolerance to stresses, particularly abiotic stress. Traditionally, the biostimulants applied to this horticultural crop are based on natural products (plant extracts, algae, microorganisms, and mineral nutrients) (Figure 6). These bioproducts have a direct effect on the plant, to stimulate its processes and favor its metabolism. These processes include nutrient absorption, increased biomass, improved fruit quality, and tolerance to biotic and abiotic stresses [64,66,67]. Other benefits include the detoxification of heavy metals, the stimulation of natural toxins for pest and disease control and the more efficient use of water [68].

4.2. Effects of Biostimulants on Morphophysiological Processes

Biostimulants optimize physiological processes in strawberry plants (Figure 7, Table 2). They activate important molecules in the plant physiological process and ensure that they are translocated to specific parts, which results in evident phenotypic changes. For example, molecules important in growth are activated by seaweed-based biostimulants, which provide macro and microelements, amino acids, vitamins, complex polysaccharides, cytokinins, auxins, and auxin-like compounds, which affect plant cell metabolism [69]. An example is the application of 5-aminolevulinic acid (natural growth regulator) in strawberry plants under osmotic stress, which benefited the gas exchange of photosynthesis and therefore improved the growth and development of roots exposed to severe osmotic stress [19]. According to the same authors, application of these products also helps maintain osmotic root balance in stressed plants by inducing aquaporins (PIP2 genes), regulating substances involved in absorption (such as jasmonates) and transporting and distributing Na+, as well as enabling greater absorption of water from the substrate through antioxidant enzymatic activities.
Another metabolic process involving biostimulants in strawberry plants, which is considered as secondary and is evidenced by the application of jasmonate in plants under water deficit, is the biosynthesis of terpenes (secondary metabolites that act in response to ecological stress), phenolic compounds (such as anthocyanins, flavonols and phenolic acids), isoprenoids (which play essential roles in photosynthesis, respiration, development, and defense against phytopathogens), alkaloids (which are nitrogenous compounds, carbon, oxygen, and hydrogen), and phenylpropanoids (which protect against UV light and defend against herbivores, and phytopathogens) [19].
In addition, the application of seaweed-based biostimulant (based on natural brassinolide) associated with water regimes resulted in an increase in ascorbic acid content with excessive irrigation and malic acid content with deficient irrigation [14]. The authors also showed that, under conditions of stress due to water deficit or excess, the application of brassinolide resulted in a reduction, in relation to the control (irrigation of 50% in relation to the control irrigation regime—397 mm cycle−1—without biostimulants application), in the contents of phenolic acids (ellagic acid, synapic acid, caffeic acid, syringic acid, chlorogenic acid, and p-coumaric acid), flavonoids (catechin, epigallocatechin, quercetin gallate, epicatechin, and rutin), and anthocyanins (cyanidin-3-O-glucoside, pelargonidin-3-O-glucoside, and pelargonidin-3-O-rutinoside) [14]. These compounds are associated with plant defense mechanisms under controlled stress. They are beneficial for maintaining productivity in adverse situations and have positive nutraceutical attributes due to their antioxidant activity. In this context, the results highlight the importance of applying biostimulants alongside water regimes involving mild to moderate deficits to obtain higher-quality fruit [70].
Increased nutraceutical activity of strawberries was reported after the application of salicylic acid (0.125 μM L−1), glutamic acid (5 g L−1), and cysteine (50 mg L−1) [3]. In this study, an increase of 21% in the content of total soluble solids, 3.5% in antioxidant capacity, and 59% in phenolic compounds was evidenced. In addition, it was revealed that the isolated application of salicylic acid increased phenolic compounds by 67% and vitamin C by 32%. Salicylic acid plays important roles in mitigating abiotic stresses, such as salinity, drought, cold, heat, and heavy metals [68]. The application of salicylic acid (50 mg L−1) or chitosan (50 mg L−1) has also been identified as responsible for the increase in vitamin C, soluble solids, total sugars, and anthocyanin in strawberries, even after 16 days of storage [2].
Humic and fulvic acids are also studied as biostimulants in strawberry plants. These compounds have previously been shown to increase salt stress tolerance [71]. Humic acid stimulates plant growth and yield to suppress disease and to provide greater stress resistance, while fulvic acid improves the root system, growth rate, and yield [68]. Also, the use of humic acid in strawberry plants improved the nutrient utilization of organic fertilizers [72]. In addition, the use of fulvic acid associated with Azospirillum increased the number of leaves by 38.8%, the root volume by 42.6%, the fresh mass by 130%, the dry mass by 63.8%, the number of fruits by 50%, and the yield by 59.5% [73]. In the same study, fulvic acid associated with Pseudomonas fluorescens favored photosynthesis and increased the levels of total soluble solids and vitamin C by 25% and 17.1%, respectively.
The use of the symbiotic bacterium Methylobacterium symbioticum in strawberry showed a reduction in the supply of 25% of nitrogen, concomitant with a reduction in nitrate reductase activity and an increase in photosynthetic efficiency [74]. The use of a plant extract-based biostimulant (composed of 250 ppm gibberellic acid, 1.5% K2O, 18% alginic acid, and 67% organic acid) increased the leaf contents of nitrogen, phosphorus, potassium, calcium, magnesium, and manganese [57]. The same authors state that the nutritional quality of the plant was related to water availability and the choice of cultivars. The use of entomopathogenic fungi (Beauveria bassiana and Metarhizium brunneum) has also been described as a biostimulant in strawberry plants, capable of inducing defense responses that are still unknown [75,76].
Regarding the morphological attributes, it was found that the use of biostimulants based on seaweed extract Ascophyllum nodosum associated with microbial biostimulants (Bacillus subtilis, Bacillus pumilus, Bacillus licheniformis, Bacillus megaterium, Bacillus amyloliquefaciens, and Trichoderma harzianum) modified the architecture of the plant’s root system (increase in dry mass by up to 32% and in the total surface by up to 28%), which optimized nitrogen uptake, and resulted in a greater number of crowns and greater plant respiration [77].
Table 2. Types and effects of biostimulants applied to strawberry plants described in the literature (2020–2025).
Table 2. Types and effects of biostimulants applied to strawberry plants described in the literature (2020–2025).
BiostimulantsEffectsReferences
Silicon
  • Increase in biomass and yield;
  • Improvement in plant defense mechanisms by activating antioxidant compounds and enzymes (abscisic acid, benzoic acids, and derivatives of ellagic acids).
[78,79]
Potassium silicate and potassium phosphite compound
  • It did not improve flavor, aroma, and color of fruits;
  • Increase in phenolic acids and flavonoids.
[9]
Nanoselenium
  • Reduction in pesticide use;
  • Increased antioxidant activity and soluble sugar content;
  • Reduction in fruit water loss during storage.
[80]
Zeolite, kaolin, and chitosan
  • Use of zeolite improved yield and fruit quality and nutrient content in leaves;
  • Joint applications of the three biostimulants (zeolite, kaolin, and chitosan) obtained results increasing to those observed only with zeolite, regarding the increase in yield and fruit quality, as well as in the nutritional content of the plant.
[81,82]
Seaweed extracts associated with microorganisms
  • Improvement in photosynthetic performance and increase in yield;
  • Changes in secondary compounds (ascorbic acid and malic acid).
[11,14,77]
Seaweed extracts (Durvillaea potatorum and Ascophyllum nodosum)
  • Increase in leaf area and number of crowns per plant;
  • Better yield per plant and increased fruit conservation potential;
  • Higher contents of total soluble solids and anthocyanins in fruits;
  • Increased concentrations of hydrogen peroxide in cells.
[69,83]
Ascophyllum nodosum
  • There were no effects on vegetative growth;
  • Improvement in photosynthetic rate, even in plants under water stress;
  • In plants without water stress there was an improvement in fruit quality.
[70,84]
Ecklonia maxima
  • Stimulation of plant growth, yield, and nutritive and nutraceutical characteristics of fruits;
  • In association with iodine, the concentration of ascorbic acid, phenol, and anthocyanins was increased.
[85]
Scenedesmus sp.
  • Accumulation of hormonal analogs of auxin and cytokinin, promoting plant growth;
  • It affected the microbiota of the strawberry tree (increase in taxa Actinospica and Streptomyces).
[86]
Thymus capitatus essential oil and microalgae consortium (Chlorella sp., Scenedesmus sp., Spirulina sp., and Synechocystis sp.)
  • Increase of 74% in the radical system, 21.1% in the number of leaves, and 24.3% in the aerial part;
  • Improvement in chlorophyll content;
  • Increase in peroxidase activity and phenol and hydrogen peroxide contents in leaves.
[87]
Amino acids and yeast extracts
  • Crown development, above-ground biomass accumulation was increased and higher fruit production.
[88]
Polyamines
  • There are aliphatic amines naturally present in plants; among them, putrescine, spermidine, and spermine are the main polyamines found in plants, being related to processes of proliferation, growth, morphogenesis, and cell differentiation;
  • The exogenous application of polyamines can be a promising alternative to increase plant tolerance to stress.
[89]
Humic acid
  • The dose of 5 L ha−1 and associated with organic fertilizer increased plant growth and fruit yield and quality.
[72]
Fulvic acid
  • With Azospirillum the number of leaves, radial volume, biomass, and productivity increased;
  • With Pseudomonas fluorescens it improved photosynthesis and the sugar and vitamin C contents of the fruits.
[73]
Salicylic acid, glutamic acid, and cysteine
  • Associated, increased fruit weight, and diameter, in addition to improving nutraceutical quality;
  • Salicylic acid associated with chitosan increased soluble solids and anthocyanin in fruits;
  • Plant-associated glutamic acid was able to modify soil microbiota.
[2,3,90]
Citric acid
  • Increase in the number of flower stems and shoot biomass;
  • Increase in calcium and leaf nitrogen content;
  • Increase in yield.
[63]
Fermented kiwi
  • Improvement in physiological and morphological attributes and fruit quality.
[91]
Hydroalcoholic extracts of Calendula officinalis, Salvia officinalis, Tagetes sp., and Taraxacum officinalis
  • Increased biomass and root system;
  • C. officinalis improved the root architecture.
[92]
Extract of Moringa oleifera L.
  • Nutrient supply, regulation of hormones and amino acids;
  • Improved fruit quality.
[93]
Trichoderma atroviride and vegetable protein hydrolysate
  • In simultaneous application, there was an increase in commercial yield, average fruit weight, fruit dry matter, total sugars, ascorbic acid, flavonoids, anthocyanins, total polyphenols, and antioxidant activity.
[94]
Trichoderma harzianum strain T22
  • Increase in the contents of sugars, total polyphenols and anthocyanins in fruits.
[63]
Azospirillum brasilense, Gluconacetobacter diazotrophicus, and Bacillus amyloliquefaciens
  • Increase in the number of flower stems;
  • Increase in crown diameter;
  • Increase in dry biomass and shoot flask;
  • Increase in leaf nitrogen content;
  • Combined with Rhizoglomus irregularis there was an increase in flower stems and sugar content.
[63]
Bacillus pumilis and Ampelomyces
  • Powdery mildew control, with integrated use of biostimulants, biopesticides, and chemical fungicides.
[95]
Arthrobacter agilis UMCV2 and Bacillus methylotrophicus M4-96
  • Different mechanisms of plant biostimulation (emission of volatile and diffusible compounds);
  • They can be inoculated in micropropagated plants and persist in colonized plant tissues;
  • Increased yield and quality of fruits in greenhouse conditions.
[96]
Methylobacterium symbioticum
  • Nitrogen supply to the plant.
[74]
Beauveria bassiana
  • Induction of systemic responses in the plant that affected populations of thrips and aphids (by interrupting reproduction and causing feeding disturbances).
[75]
Metarhizium brunneum
  • It generates direct effects (regulation of hormones) and indirect effects (protective properties);
  • Need for further studies to describe the physiology and chemical ecology of this interaction.
[76]

4.3. State-of-the-Art on Biostimulants as Mitigators of Water Deficit in Strawberry Plants

Water stress causes morphophysiological changes, yield, and fruit quality of strawberry plants. Therefore, limitations in plant growth and development are described, and methods are needed to induce resistance and protect plants from this stressor phenomenon [97]. In addition to technical approaches, water-saving strategies are necessary, such as the establishment of cultivars that require low water consumption and are capable of maintaining productivity [30]. Another way to mitigate water stress in strawberry is the use of biostimulants, such as inoculation with beneficial microorganisms in the rhizosphere of plants [97]. In addition to microbial agents, there are also non-microbial biostimulants, which can be used as a strategy to mitigate plant stresses [65].
Biostimulants can increase the antioxidant activity and defense mechanisms of the plant by activating enzymes, plant regulators, and antioxidant compounds, such as hydrogen peroxide [83,86,87], or even by the organization of cell walls [78]. Therefore, biostimulants represent a tool not only with the potential to increase strawberry yield and quality, but also because they can help in resilience to biotic and abiotic stresses. However, the physiological and biochemical processes involved in the improvements achieved with the use of biostimulants in strawberry still need to be better understood and require further research [64,75,76,91].
It is already known that effective cultivation practices, such as choosing suitable cultivars, effective nutrition, and using biostimulants, can improve fruit quality [57]. Recent studies have associated the use of biostimulants with positive perspectives regarding abiotic stresses, such as water stress [98]. This sustainable management in stressful environments can improve gas exchange, so as to cause cuticle thickening (modulation of water balance), reduce electrolyte leakage, and increase levels of photosynthetic pigments, phenolic compounds, and antioxidant activity [70,78]. This is because problems with limited water resources have affected social and agricultural activities on a global scale [3].
We emphasize the variability in results of biostimulants in relation to strawberry cultivation systems [63]. These differences may be due to the properties of the growing medium (organic or conventional) and climatic factors (light, temperature, and water resources). In view of this data asymmetry, there is a need for research on the use of biostimulants in greenhouse plants, as well as those submitted to different abiotic conditions. This will optimize the applicability of these bioproducts from the determination of doses, frequencies, and application methods.
Another research gap that remains open is related to the study of the economic feasibility and sustainability of integrating the use of biostimulants in strawberry cultivation [63,88]. This is because, despite the benefits of increased growth, yield, and resilience of the plant to stress, commercial products in this sector remain limited, as well as information about their production and applicability. In addition, to meet the demands of biostimulants from agriculture, from a commercial point of view, production is dependent on the challenging transfer of technology to an industrial scale. Therefore, the commercial scalability of biostimulant production requires technical-economic analysis and search for optimal production parameters to induce the production of bioactive compounds and improve the enzymatic activity, that meet the expectations of the producer who seeks to enhance the fruit quality [99]. Therefore, in addition to studies on the applicability of biostimulants, studies on the standardization of protocols are also needed [82], from production optimization and standardization to the commercialization of these products.
As for the water regimes used in the cultivation of strawberry in greenhouses, it is known that excessive irrigation, in addition to being divergent from an ecological point of view, can be harmful to the morphophysiology and fruit quality and facilitate the proliferation of pathogens. Studies have explored deficient irrigation systems in the cultivation of this horticultural crop. The main positive effect is to improve the efficiency of water use, protect the environment, and increase yield and fruit quality [37]. However, the critical limit, which generates a balance between strawberry growth and water consumption, has not yet been easily quantified [48].
Studies that focus on strawberry irrigation systems, with ‘Romina’, ‘Sibilla’, ‘Cristina’, ‘Albion’, ‘San Andreas’, and ‘Monterey’ cultivars, indicate that with a 20% reduction in the total irrigated water (irrigation rate of 957 m3 ha−1), results similar to the control (100% irrigation—irrigation rate of 1183 m3 ha−1) were obtained in terms of fruit firmness, sugar content, folate content, and antioxidant capacity, without reducing yield and ensuring savings of up to 226 m3 of water per hectare and crop cycle, in an experiment conducted in open fields, with strawberry plants grown in soil and covered by a plastic tunnel [98]. On the other hand, taking into account the production of flowers and fruits, the literature indicates that irrigation maintaining 75% of the field capacity (mild water deficit) showed the best result in yield and optimization of water use by strawberries in an experiment conducted in a greenhouse with plants grown in pots filled with soil, cocopeat, charcoal husk and goat manure (3:2:2:1, v/v/v/v) mixture [34]. Other results demonstrate that intermittent sprinkler irrigation made it possible to maintain yield and reduced water consumption by 50 to 67% during the establishment of bare-rooted strawberry daughter plants, representing a water saving of 322 to 429 mm (3.2 to 4.3 million liters per hectare) [45].
The genetic and physiological mechanisms of how deficient irrigation affects strawberry still need to be better understood to describe the modulation to produce substances related to oxidative stress as defense mechanisms, in addition to knowing how these changes can affect yield and fruit quality. Therefore, new irrigation strategies are needed to increase the resilience of food production, reduce negative environmental impacts, and promote economic and agricultural sustainability [37]. In this aspect, in the search for the economy of natural resources aiming at places of low availability of surface water [100], the selection of cultivars with lower water requirement, and high photosynthetic activity [20] represents an interesting strategy [38,60].
Therefore, to mitigate water stress, it is essential to implement sustainable practices, precision irrigation, sensing through terrestrial devices and robotics (for real-time data provision), as well as developing drought-resistant cultivars [18,101]. Effective and innovative solutions, such as the use of biostimulants and adequate irrigation, are crucial to optimize water use in intensive crops, such as strawberry.

5. Final Considerations and Future Perspectives

Agriculture, representing the integration of natural resources and human activity, is one of the sectors that consumes the most water in the world. However, increasing water scarcity threatens this ancient practice. Along with climate change, resulting from global warming, there is a deregulation in the rainfall regime, which results in the limitation of water resources, previously considered unlimited. Seeking to overcome these challenges, models for predicting climate variables still need improvement, considering a water scenario planning [102]. Solutions considering the reconstruction of natural resources bring with them social, cultural, and economic aspects, highlighting a circular economy through robust proposals that consider the biodiversity preservation [103]. Therefore, the search for tools that maintain or increase food productivity and quality, despite the scarcity of water resources, is already part of research around the world, requiring wisdom and innovation. The search for healthy food in sufficient quantity is no longer a demand of the future, but an urgency of the present. Currently, around 3 billion people worldwide do not have access to a healthy diet, and the outlook is for an increase of up to 70% in food demand by 2050 [104,105].
Among these foods, strawberries are already consolidated for their sensory and nutritional acceptance. However, their production still suffers from the major consequences of climate stress, particularly water deficits. Therefore, irrigation management must follow a fine adjustment, since excessive irrigation, in addition to intensifying the production of more acidic fruits, results in water waste [98,106]. On the other hand, deficient irrigation systems, although they produce better quality fruit, can compromise yield and impact economic viability. Under stress conditions, plants, in the face of stress, in addition to reducing energy obtainment, overload their oxidative mechanisms [107,108].
To address these agricultural challenges, tools such as plant improvement, morphophysiological studies, management to induce stress tolerance (such as biostimulation), and the use of artificial intelligence technologies (which promise to expand our ability to understand and intervene accurately) are necessary to mitigate plant stress.
The emergence of a market for biostimulants, founded on solid and ethical principles, presents an alternative approach for productions seeking to reduce water consumption while adhering to sustainability criteria. This is because, for the most part, biostimulants are of natural origin and without the addition of synthetics and, when certified, can be used in organic agriculture. Despite their intensive use in organic farming, biostimulants are also applied in conventional farming, especially in agroecological transition systems, aiming to reduce the environmental impacts caused by biocides. In addition to acting in synchrony with other beneficial microorganisms [109], biostimulants act in the plant’s defense through the supplementation of hormones and other compounds, being a source of intermediate molecules for defense against oxidative stress [110]. Thus, it is important to highlight the formation of a genotype × environment × management × biostimulant system [111].
The long-term effects of such practices, including the procurement of quality food that respects the ethical premises of sustainability and ensures the prosperity of the producer, are of paramount importance. Biostimulants represent merely one of the tools at the disposal of those engaged in agricultural practice. The effectiveness of this particular tool is the result of a harmonious integration of numerous elements of knowledge and agents, including producers, researchers and public authorities. It is expected that robust practices will be implemented to ensure the health benefits and profitability of production.

Author Contributions

Conceptualization, J.L.C., M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; methodology, J.L.C., L.V.C.M. and J.L.T.C.; validation, J.L.C., L.V.C.M., M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; formal analysis, J.L.C., L.V.C.M. and J.L.T.C.; investigation, J.L.C., M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; resources, M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; data curation, J.L.C. and L.V.C.M.; writing—original draft preparation, J.L.C., M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; writing—review and editing, J.L.C., M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; visualization, J.L.C., L.V.C.M., M.P.B., L.M.C., F.W.R.J., P.P. and J.L.T.C.; supervision, M.P.B., P.P. and J.L.T.C.; project administration, J.L.T.C.; funding acquisition, P.P. and J.L.T.C. All authors have read and agreed to the published version of the manuscript.

Funding

This study was financed by the Coordination for the Improvement of Higher Education Personnel (CAPES) (finance code 001).

Data Availability Statement

Not applicable.

Acknowledgments

To University of Passo Fundo and CAPES.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Bibliometric research on the use of biostimulants in strawberry plants under water stress.
Figure 1. Bibliometric research on the use of biostimulants in strawberry plants under water stress.
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Figure 2. Number of studies on the use of biostimulants in strawberry plants under water stress, from 2020 to 2025, on the Scopus platform.
Figure 2. Number of studies on the use of biostimulants in strawberry plants under water stress, from 2020 to 2025, on the Scopus platform.
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Figure 3. Density of publications for the bibliometric search of “fragaria” OR “strawberry” AND “biostimulant” (A) and the search for “fragaria” OR “strawberry” AND “water stress” (B).
Figure 3. Density of publications for the bibliometric search of “fragaria” OR “strawberry” AND “biostimulant” (A) and the search for “fragaria” OR “strawberry” AND “water stress” (B).
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Figure 4. Word cloud of the bibliometric search of “fragaria” OR “strawberry” AND “biostimulant” (A) and of the search of “fragaria” OR “strawberry” AND “water stress” (B). In the first case (A), the yellow cluster associated biostimulation with fruit quality, the green cluster combined yield and plant growth with fruit quality, the red cluster associated Fragaria with biostimulation, and the blue cluster related abiotic stresses with chitosan. In the second case (B), the green cluster associated Fragaria with chlorophyll and botany, the red cluster combined strawberry water stress with productivity and growth, and the blue cluster associated water stress with controlled studies and photosynthesis.
Figure 4. Word cloud of the bibliometric search of “fragaria” OR “strawberry” AND “biostimulant” (A) and of the search of “fragaria” OR “strawberry” AND “water stress” (B). In the first case (A), the yellow cluster associated biostimulation with fruit quality, the green cluster combined yield and plant growth with fruit quality, the red cluster associated Fragaria with biostimulation, and the blue cluster related abiotic stresses with chitosan. In the second case (B), the green cluster associated Fragaria with chlorophyll and botany, the red cluster combined strawberry water stress with productivity and growth, and the blue cluster associated water stress with controlled studies and photosynthesis.
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Figure 5. Irrigation intensities in strawberry—described in the literature as excessive (125%) [35], normal (100%) [36], and mild (75%) [37], moderate (50%) [25,38], and intense (25%) [25] deficits—associated with morphophysiological and fruit quality responses. This association indicates the need for an adjustment regarding the intensity of irrigation (seeking to save water resources) and the maintenance of productivity and quality of fruits.
Figure 5. Irrigation intensities in strawberry—described in the literature as excessive (125%) [35], normal (100%) [36], and mild (75%) [37], moderate (50%) [25,38], and intense (25%) [25] deficits—associated with morphophysiological and fruit quality responses. This association indicates the need for an adjustment regarding the intensity of irrigation (seeking to save water resources) and the maintenance of productivity and quality of fruits.
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Figure 6. Graphic scheme of the requirements for use and types of biostimulants applied to strawberry.
Figure 6. Graphic scheme of the requirements for use and types of biostimulants applied to strawberry.
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Figure 7. Performance of biostimulants in strawberry organs. The arrows indicate increases in the attributes described.
Figure 7. Performance of biostimulants in strawberry organs. The arrows indicate increases in the attributes described.
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MDPI and ACS Style

Cassel, J.L.; Maldaner, L.V.C.; Bortoluzzi, M.P.; Colla, L.M.; Reichert Junior, F.W.; Palencia, P.; Chiomento, J.L.T. Biostimulants as a Tool for Mitigating Water Deficit Stress in Strawberry Cultivation. Agronomy 2025, 15, 2643. https://doi.org/10.3390/agronomy15112643

AMA Style

Cassel JL, Maldaner LVC, Bortoluzzi MP, Colla LM, Reichert Junior FW, Palencia P, Chiomento JLT. Biostimulants as a Tool for Mitigating Water Deficit Stress in Strawberry Cultivation. Agronomy. 2025; 15(11):2643. https://doi.org/10.3390/agronomy15112643

Chicago/Turabian Style

Cassel, Júlia Letícia, Laura Valentina Caus Maldaner, Mateus Possebon Bortoluzzi, Luciane Maria Colla, Francisco Wilson Reichert Junior, Pedro Palencia, and José Luís Trevizan Chiomento. 2025. "Biostimulants as a Tool for Mitigating Water Deficit Stress in Strawberry Cultivation" Agronomy 15, no. 11: 2643. https://doi.org/10.3390/agronomy15112643

APA Style

Cassel, J. L., Maldaner, L. V. C., Bortoluzzi, M. P., Colla, L. M., Reichert Junior, F. W., Palencia, P., & Chiomento, J. L. T. (2025). Biostimulants as a Tool for Mitigating Water Deficit Stress in Strawberry Cultivation. Agronomy, 15(11), 2643. https://doi.org/10.3390/agronomy15112643

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