Adrian Alejandro Espinosa-Antón1 Eduardo Salcedo-Perez1*
, Rosalba Mireya Hernández-Herrera1
1 University Center of Biological and Agricultural Sciences, University of Guadalajara. Camino Ramón Padilla Sánchez No. 2100, C.P. 45200, Zapopan, Jalisco, Mexico.
Extractos botánicos con potencial bioestimulante para la agricultura en México: una revisión
Articulo de revisión
The main challenge facing the agricultural sector today is to ensure continued access to enough nutritious food with minimal environmental impact. Therefore, the search for sustainable and effective methods to stimulate crop yield and quality is a priority. Plants are emerging as an attractive local resource for biostimulant extracts due to their abun- dant biomass and high content of bioactive compounds. However, the factors that support the implementation of botanical extracts as alternative bioproducts to reduce de- pendence on agrochemicals are not yet fully understood. This review aimed to analyze the available information on the potential use of botanical extracts as biostimulants for Mexican agriculture. An updated overview of the definition and regulation of agricultural biostimulants worldwide and in Mexico is provided. In addition, research trends on the biostimulant properties of extracts derived from Mexican plants related to the species used as raw material, extraction methods, modes of application, and types of crops evaluated are analyzed. Finally, the effects and mechanisms of action of botanical extracts on crop plants are discussed, and future research needs to optimize their application in agricultural production.
El reto principal del sector agrícola actualmente es garantizar el acceso constante a cantidades suficientes de alimentos nutritivos con un impacto ambiental mínimo. Por ende, la búsqueda de métodos sostenibles y eficaces para estimular el rendimiento y calidad de las cosechas representa una prioridad. Las plantas son recursos locales atractivos para la obtención de extractos bioestimulantes debido a su disponibilidad de biomasa y alto contenido en compuestos bioactivos. Sin embargo, la comprensión de los factores que sustentan la aplicación de los extractos botánicos como bioproductos alternativos para reducir la dependencia de los agroquímicos es aún limitada. El objetivo de esta revisión es analizar la información disponible sobre el uso potencial de los extractos botánicos como bioestimulantes para la agri- cultura mexicana. Se ofrece una visión general y actualizada
*Author for correspondence: Eduardo Salcedo-Perez
e-mail: eduardo.salcedo@academicos.udg.mx, pizanjc@gmail.com
Received: April 9, 2025
Accepted: May 12, 2025
Published: June 13, 2025
de la definición y regulación de los bioestimulantes agrícolas en el mundo y en México. Además, se analizan las tendencias de la investigación sobre las propiedades bioestimulantes de los extractos derivados de plantas mexicanas relacionadas con las especies empleadas como materia prima, los méto- dos de extracción, las vías de aplicación y los tipos de cultivos evaluados. Finalmente, se discuten los efectos y mecanismos de acción de los extractos botánicos en los cultivos, así como las necesidades futuras de investigación para optimizar su implementación en la producción agrícola.
The production and productivity of the Mexican agricultural sector depend, among other factors, on the use of agroche- micals such as fertilizers and pesticides (DataMéxico, 2025). According to the 2019 National Agricultural Survey (ENA)
67.4 % of the sector’s production units used chemical fertili- zers, and the remaining 24.4 % used natural fertilizers (INEGI, 2019). In Mexico, as in the rest of the world, the use of fer- tilizers is mainly concentrated in nitrogenous ones (65.3 %), followed by phosphorous fertilizers (16.1 %) and potassium fertilizers (18.6 %) (Banco de México, 2022).
For decades, the dependence on synthetic chemical inputs has significantly boosted food production. However, the inadequate application of these products has also gene- rated harmful effects on human health and agroecosystems due to their high toxicity, rapid mobility in the soil, and high residuality in harvested products (Rouphael and Colla, 2020; Carvalho et al., 2021). Therefore, one of the fundamental challenges currently facing the agricultural sector is the implementation of new and efficient strategies that promote the sustainability of agroproductive systems (Ali et al., 2021; Martínez-Lorente et al., 2024). Also, the accelerated growth of the population has promoted the need to enhance the development and yield of crops to meet consumer demands in terms of quantity and quality (Zulfiqar et al., 2020; Akter et al., 2024).
In the search for sustainable solutions in agriculture, natural products derived from renewable raw materials
Volume XXVII
DOI: 10.18633/biotecnia.v27.2633
Journal of biological and health sciences http://biotecnia.unison.mx
Universidad de Sonora
ISSN: 1665-1456
emerge as a promising alternative (Jasso de Rodríguez et al., 2020; Godlewska et al., 2021). Plant biostimulants comprise substances, compounds, and growth-promoting microorga- nisms that regulate or enhance plant-soil system processes (Calvo et al., 2014; Rouphael and Colla, 2020; Sanjuán et al., 2023). Nevertheless, despite the remarkable advances in biostimulant science in recent years, their definition, classi- fication, and regulation continue to evolve. The importance of these bioproducts lies in their ability to enhance nutrient use efficiency and promote plant vigor, independently of their nutrient content and without directly affecting pest or disease control (Yakhin et al., 2017; Corsi et al., 2022). Therefo- re, the integration of biostimulant products into agricultural production systems has been demonstrated to promote desirable agronomic traits and help reduce the use of agro- chemical inputs (Caradonia et al., 2019; Han et al., 2024).
However, to generalize the implementation of biosti- mulants in agriculture, it is important to consider the avai- lability and accessibility of raw materials for processing (Xu and Geelen, 2018). In this context, there is growing interest in exploring the potential of botanical extracts as sources for novel biostimulant formulations. This interest stems from their high content of bioactive compounds, including polyphenols, phytohormones, organic acids, sterols, and terpenoids, as well as their availability from diverse and abundant raw materials, including weeds, invasive species, and agro-industrial byproducts (Godlewska et al., 2021; Han et al., 2024). Moreover, the cost-effectiveness of their produc- tion by the same manufacturer underscores their economic viability (Ahmad et al., 2022; Velasco-Ramírez et al., 2022b).
Due to the novelty of this topic, understanding the fac- tors that underpin the use of botanical extracts as alternative bioproducts, to improve agricultural production and reduce dependence on agrochemicals, is still limited. The main objective of the current review was to analyze the available scientific information on the potential use of botanical ex- tracts as biostimulants for Mexican agriculture. In particular, this review provides an updated overview of the definition and regulation of agricultural biostimulants worldwide and in Mexico. Furthermore, it examines current research trends on botanical extracts, focusing on the species used as raw materials, extraction methods, modes of application, and the types of crops evaluated. Finally, the effects and mechanisms of action of botanical extracts on plants are discussed, along with future research needs to optimize their application in agricultural production.
Global and Mexican context of plant biostimulants Although interest in biostimulants in agriculture began in the second half of the twentieth century, in recent years it has experience exponential growth, consolidating itself as an innovative tool to improve the production and productivity of crops (Ahmad et al., 2022; Corsi et al., 2022; Han et al., 2024). Throughout history, plant biostimulants have received various names in the scientific literature, such as biogenic stimulants, plant strengtheners, metabolic enhancers,
phytostimulants, positive plant growth regulators, growth promoters and biofertilizers, among others (Yakhin et al., 2017). However, there is no single, universally accepted definition of the term “plant biostimulant” (Sanjuán et al., 2023).
This lack of consensus is due to multiple factors, among which the following stand out: (1) heterogeneity in their sources of production, elaboration processes, and compo- sition; (2) the difficulty in elucidating its mode of action; (3) similarities with other categories of agricultural products regulated under existing regulations; (4) the variability in re- gulatory frameworks between countries and regions; (5) the commercial interests of large companies that dominate the agrochemical market and; (6) the progressive advancement of scientific knowledge about the nature of its effects on the different components of the plant-soil systems (Yakhin et al., 2017; Corsi et al., 2022; Sanjuán et al., 2024).
Despite this context, the academic community agrees that biostimulants are substances and microorganisms that, when applied at low concentrations to plants, seeds, or the rhizosphere, stimulate natural processes that improve or benefit nutrient absorption, nutritional efficiency, tolerance to abiotic stress, growth, productivity, and/or crop quality regardless of their nutrient content (du Jardin, 2015; Yakhin et al., 2017; Zulfiqar et al., 2019; Han et al., 2024; Sanjuán et al., 2024). By extension, products containing mixtures of such substances and/or microorganisms are also recognized as plant biostimulants (du Jardin, 2015).
One of the most supported premises is that the recog- nition of a product as a plant biostimulant should focus on its agricultural functions, rather than on the nature of its components or its modes of action, as these can originate from multiple sources (Ricci et al., 2019; Sanjuán et al., 2024). These include humic substances, animal and plant protein hydrolysates, algae and plant extracts, biochemicals, bene- ficial microorganisms, and agroindustrial residues (Yakhin et al., 2017). More recently, a trend has been to classify this wide diversity into microbial and non-microbial biostimulants (Corsi et al., 2022).
Commercial biostimulants may include compounds not naturally present in the feedstock (Yakhin et al., 2017; Sanjuán et al., 2024). Adding to the lack of a clear definition and con- sistent criteria that distinguish biostimulants, this situation generates confusion with other agricultural formulations such as fertilizers and growth regulators (Corsi et al., 2022). To address this problem, it is essential to establish a regulatory framework that guarantees the quality, efficacy, and safety of biostimulants at the national and regional levels (Yakhin et al., 2017; Ricci et al., 2019).
In this sense, the European Union established the “Re- gulation (EU) 2019/1009 of the European Parliament and the Council”, which is the first legislative text that provides a legal framework for the marketing of biostimulant products (European Union, 2019). On the other hand, in the United States, despite significant progress in defining and proposing a bill with the backing of regulatory agencies and industry, there is still no federal or state regulation for these products
(GPO, 2023). There are no specific legal provisions for plant biostimulants in other countries such as India, South Africa, Canada, and Brazil. Instead, these products are indirectly regulated under national laws that address other agricultural inputs (Caradonia et al., 2019).
In Mexico, despite the wide availability of both natio- nal and imported biostimulants in the agricultural market, current legislation lacks specific regulations for their produc- tion, labeling, and commercialization. In the absence of a le- gal definition of “plant biostimulant”, they are intended to be regulated under other categories recognized in the Norma Oficial Mexicana NOM-077-FITO-2000, which establishes the specifications, criteria, and procedures to regulate studies of the biological effectiveness of plant nutrition inputs for agricultural use (DOF, 2020). Biological effectiveness results from applying a plant nutrition input, which can be said to have a biostimulant effect if it improves nutritional uptake efficiency, abiotic stress tolerance, and/or crop quality (DOF, 2020). These inputs are highly diverse and include:
Inoculant: Plant nutrition input made from microor- ganisms that, when applied, favors the use of nutrients in association with the plant or its rhizosphere.
Organic soil improvers: plant nutrition inputs made from organic products or microorganisms that favorably modify the physical, chemical, or biological conditions of the soil for the best plant development.
Type 1 growth regulators: Growth regulators are based on substances found naturally in plant tissues, whether obtai- ned by extraction, fermentation, synthesis, or other methods. These types of regulators are also known as phytohormones or plant hormones.
Type 2 growth regulators: Products based on substances obtained by synthesis and not found naturally in the plant.
However, these definitions do not fully capture the diverse and multifunctional nature of plant biostimulants. In addition, when analyzing these definitions, it can be observed that establishing an exclusive regulation for bios- timulants would be complicated due to the overlaps in their composition and modes of action with some existing cate- gories. For example, botanical or algae extracts are complex mixtures of different bioactive compounds, including plant hormones and their precursors or intermediates. This raises the question of whether they should be classified as type 1 growth regulators since they contain phytohormones, or if they should be considered biostimulants as they include substances other than phytohormones. Therefore, fostering a clear and accurate understanding of plant biostimulants is crucial to promote their adoption as sustainable alternatives to agrochemicals. Addressing this challenge requires coordi- nated efforts among academic institutions, industry stake- holders, and regulatory agencies to update and harmonize existing legislation.
In México, just over 25,000 species of vascular plants have been documented, representing approximately 9.1 % of all
known species worldwide. These figures position the country among the five nations with the most incredible wealth of vascular plants (SEMARNAT, 2012). Therefore, one of the pro- mising strategies to promote economic and environmental sustainability in Mexican agriculture is using local plant spe- cies as a source of bioactive compounds with biostimulant activity.
From 2008 to date, 15 original articles by Mexican authors have been published evaluating the use of botanical extracts as plant biostimulants. Of the total scientific production analyzed, 73 % corresponds to the years 2022 (14 %), 2023
(36 %) and 2024 (27 %). In contrast, between 2008 and 2021, the number of publications on this topic remained constant (Figure 1). The increase observed in recent years reflects an increased visibility of research that uses extracts obtained from plants as a source of biologically active substances to enhance crop growth, development, and productivity.
This growing interest in the topic can be attributed to the remarkable global boom in plant biostimulants over the past decade, which have established themselves as innova- tive alternatives to synthetic chemical fertilizers to address the challenges of sustainable agriculture (Zulfiqar et al., 2019; Ahmad et al., 2022; Han et al., 2024). Mexico is among the 10 countries with the highest production of scientific articles regarding the benefits of the use of biostimulants in agri- cultural production systems (Corsi et al., 2022), with special emphasis on research related to botanical extracts (Carvalho et al., 2022).
Also, nine scientific journals were identified that publis- hed articles included in this review, predominantly those edited by Mexican institutions. Among the Mexican journals, Biotecnia and Ecosistemas y Recursos Agropecuarios stands out for concentrating 53 % of the scientific production on the potential of plant extracts as biostimulants in agriculture. Both journals are included in the Emerging Sources Citation Index (ESCI), which is part of the primary collection of the Web of Science, and the SCIELO Citation Index, another database integrated into this same scientific information platform.
The research analyzed highlights the biostimulant properties of extracts from 22 plant species originating from diverse habitats, including semiarid zones, disturbed vegetation, halophytic grasslands, and freshwater ecosystems. These species represent approximately 0.1 % of the vascular plant diversity reported in Mexico (SEMARNAT, 2012). The analyzed species belong to 14 botanical families, with Asteraceae and Anacardaceae being the most represented, accounting for 23
% and 18 % of the total species, respectively (Figure 2A).
On the other hand, Rhus muelleri Standl. and F.A. Barkley is positioned as the most researched species as a source of extracts with biostimulant properties in agriculture, re- presenting 27 % of the articles analyzed (Figure 2B). Other prominent species include Flourensia microphylla S.F. Blake,
Figure 1. Articles published by Mexican authors on the use of botanical extracts as biostimulants for agriculture. Data for the first months of 2025 are not represented.
Figura 1. Artículos publicados por autores mexicanos sobre el uso de extractos botánicos como bioestimulantes en agricultura. Datos para los primeros meses de 2025 no están representados.
Flourensia retinophylla S.F. Blake and Verbesina sphaerocepha- la A. Gray ex S.Wats., which were the subject of study in 20 % of the studies (Figure 2B). Rhus virens Lindh. ex A. Gray and Moringa oleifera Lam. were represented in 13 % of the studies (Figure 2B). The remaining set of species assessed includes Cenchrus echinatus L., Distichlis spicata L., Rhus trilobata Nutt., Amphipterygium glaucum (Hemsl. & Rose) Hemsl. & Rose
ex Standl., Cucurbita foetidissima Kunth, Bacopa monnieri (L.) Wettst, Heliotropium curassavicum L., Flaveria trinervia (Spreng.) C. Mohr, Sesuvium verrucosum Raf., Maclura tinc- toria (L.) D. Don ex Steud, Myriophyllum aquaticum (Vell.) Verdc., Jatropha dioica Cerv., Juglans mollis Engelm., Opuntia ficus-indica (L.) Mill., and Coriandrum sativum L. Each species represents 5 % of the studies included in this work.
The biogeographical status of the plant species reveals that 59 % are native and 27 % are endemic to Mexico (DOF, 2010) (Figure 2C). The utilization of local flora as a source of bioactive extracts underscores the value of national resour- ces, and opens new possibilities for discovery of compounds that stimulate plant growth (Godlewska et al., 2021). In addi- tion, using weeds as raw material for obtaining biostimulant extracts, as in the case of C. echinatus, could represent a promising strategy (Huez-Lopez et al., 2008; Leopardi-Verde et al., 2021).
In contrast, 14 % of the species evaluated are exotic (Figure 2C), that is, those not native to Mexico and have been introduced intentionally or accidentally. Among these, M. aquaticum is classified as an invasive species (Bonilla-Barbosa and Santamaría, 2013), whose benefits in horticultural pro- duction were documented by Viveros-Legorreta et al. (2018).
Botanical extracts are complex mixtures of primary and se- condary metabolites whose biostimulant properties depend on various factors (Godlewska et al., 2021). One of the key
Figure 2. (A) Number of species evaluated by botanical family, (B) Main species evaluated as a source of biostimulant extracts, and (C) Percentage of the species studied according to their biogeographical status in México.
Figura 2. (A) Número de especies evaluadas por familia botánica, (B) Principales especies evaluadas como fuente de extractos bioestimulantes y
(C) Porcentaje de especies estudiadas de acuerdo a su estado migratorio en México.
aspects is the selection of the plant biomass source used as a feedstock, which largely determines the chemical composi- tion of the resulting extract (Zulfiqar et al., 2019; Ahmad et al., 2022). As shown in Table 1, the extracts evaluated in different studies as stimulants of crop growth and development are obtained from both whole plants and specific parts, including leaves, cladodes, or fruits. In addition, agricultural residues, often underutilized, represent valuable sources for obtaining plant-based biostimulants (Huez-López et al., 2008).
Once the plant material has been selected, several biomass pretreatment steps must be carried out before extracting the bioactive compounds from the plant tissues (Godlewska et al., 2021). The first step is to wash the collec- ted material several times with tap water to remove surface impurities (Ahmad et al., 2022). The biomass intended for extraction can then be used in its fresh state or subjected to drying processes (Table 1). These processes can be carried out at room temperature (~26 °C) in the laboratory (Velasco- Ramírez et al., 2021; 2022a) or at low temperatures (-80 °C) by freeze-drying to preserve the properties of the compounds present in the extract (e.g., polyphenols and flavonoids) (Garza-García et al., 2023). The dried plant material is then crushed and screened to obtain a homogeneous sample and increase the surface-to-volume ratio (Michalak and Chojnacka, 2014). In most studies, the crushing stage is not described in detail, although it is usually done with the help of an electric mill or mixer.
Several methods exist to obtain biostimulant extracts from plant biomass (Godlewska et al., 2021; Han et al., 2024). However, the method and solvent selection are essential to ensure a high yield of the bioactive compounds of interest and preserve their integrity and properties (Michalak and Chojnacka, 2014; Ahmad et al., 2022). In addition, identi- fying an economical, simple, effective, and environmentally friendly method for preparing biostimulants based on plant extracts is crucial to scaling up the process (Michalak and Chojnacka, 2014). The research analyzed in this study descri- bes methods for extracting plant biomass that use various approaches (Table 1).
The 54 % of the studies used physical methods for the extraction of bioactive compounds, such as maceration, au- toclaving, or ultrasound, which facilitates the release of these from plant tissue (Velasco-Ramírez et al., 2022a; Garza-García et al., 2023; Moreno-Guerrero et al., 2024; Velazquez-Gamboa et al., 2024). The choice of solvent is crucial for extraction efficiency (Michalak and Chojnacka, 2014). The 38 % of the research used organic solvents such as methanol, ethanol, and acetone, which selectively dissolve the compounds of interest (Jasso de Rodríguez et al., 2020; Brazales-Cevallos et al., 2022). The evaporation of these solvents ensures that the final extract is safe for crops application (Godlewska et al., 2021; Ahmad et al., 2022).
In addition to the methods previously described, 15
% of the studies included in this work employed biological processes such as fermentation to obtain plant extracts with biostimulant activity. This method consists in inoculating fresh plant material with natural sources of microorganisms, such as manure, which are responsible for carrying out the fermentation process under anaerobic conditions (Huez-Ló- pez et al., 2008; Alcalán-López et al., 2023). Microbial activity facilitates the release of bioactive compounds in the plant matrix and improves its chemical and functional properties, increasing its effectiveness in agricultural applications (Pérez- Alva et al., 2022).
In general, water-based extraction methods are the most common to produce botanical extracts for use in agri- culture (Table 1). These extractions are simple and relatively inexpensive and dispense with the use of organic solvents, reducing toxicity and residue generation while improving crop yield and quality (Godlewska et al., 2021; Velasco- Ramírez et al., 2021). However, using water as a solvent for specific compound extractions has disadvantages, as its low selectivity and extraction efficiency reduce the yield per unit of biomass used as feedstock (Michalak and Chojnacka, 2014).
The application method of botanical extracts determines their effectiveness as agricultural biostimulants (Han et al.,
Table 1. Summary of the methods for obtaining plant extracts as biostimulants.
Tabla 1. Resumen de métodos para la obtención de extractos de plantas como bioestimulantes.
Type of extraction | Method | Solvent | Plant material | Biomass pretreatment | References |
Chemistry | Organic solvent extraction | Methanol Ethanol/ Acetone | Leaves and fruits Leaves | Drying and crushing Fresh | Jasso de Rodríguez et al. (2020; 2023; 2024a; 2024b) Brazales-Cevallos et al. (2022) |
Water | Young leaves | Drying and crushing | Velasco-Ramírez et al. (2021; 2022; 2023) | ||
Mechanical extraction with blender | Water | Leaves | Fresh | Brazales-Cevallos et al. (2022) | |
Physics | Maceration | Water Methanol | Leaves Leaves and cladodes | Drying and crushing Fresh | Garza-García et al. (2023) Moreno-Guerrero et al. (2024) |
Ultrasound-assisted extraction | Water | Leaves | Fresh | Velázquez-Gamboa et al. (2024) | |
Biological | Anaerobic fermentation | Water Water | Waste The whole plant | Fresh Fresh and segmented | Huez-Lopez et al. (2008) Alcalán-López et al. (2023) |
2024). In this study, four methods for applying plant extracts in crops were identified and evaluated under various experi- mental conditions. These methods include seed treatment, direct application to plants or soil, and a combination of the latter approaches (Figure 3).
The 47 % of the studies explored the biostimulant effects of seed treatment by hydration with different concentrations of extracts. Of these, 27 % were performed in petri dishes under laboratory conditions, 7 % in germination trays or pots under greenhouse conditions, and 13 % in both experimen- tal environments (Figure 3). Also, direct application to plants by foliar spray accounted for 47 % of the research analyzed in this study, all of which were conducted under greenhouse conditions (Figure 3).
These results highlight the potentialities of using bota- nical extracts by both application methods to promote de- sirable agronomic traits in crops. For example, seed priming with botanical extracts is consolidated as an effective pre- conditioning strategy, which optimizes germination capacity and promotes adequate seedling establishment (Alcalán- López et al., 2023; Velasco-Ramírez et al., 2022a; Carvalho et al., 2024; Jasso de Rodríguez et al., 2024b). On the other hand, foliar spraying allows an immediate interaction of the bioactive substances present in plant extracts with plant tis- sues, thanks to their rapid absorption through stomata and cuticles, which induces a direct response in crops (Youssef et al., 2022; Han et al., 2024). This application method is most effective when it is performed in the morning, between 8:00 and 10:00 a.m., when the stomata remain open (Jasso de Rodríguez et al., 2023). Likewise, the complementary use of agricultural adjuvants contributes to optimizing the absorp- tion of extracts through plant foliage (Jasso de Rodríguez et al., 2020).
The applications of botanical extracts by irrigation to the soil or to the growing substrate were evaluated in 34 % of the studies, 27 % in the greenhouse, and 7 % in the labo- ratory (Figure 3). This method delivers plant extracts to the
Figure 3. Main applications of plant extracts in agriculture, and the experimental conditions evaluated in the articles published by Mexican authors during the study period.
Figura 3. Principales aplicaciones de los extractos vegetales en agricultura, y las condiciones experimentales evaluadas en publicaciones por autores mexicanos durante el periodo de estudio.
base of the crop, promoting root development and microbial activity in the rhizosphere (Youssef et al., 2022). However, bioactive substances can be adsorbed by soil particles or lost through leaching, limiting their bioavailability (Mattner et al., 2013; Han et al., 2024). The combined application of botanical extracts to both plants and soil has been demonstrated to enhance their effectiveness (Velasco-Ramírez et al., 2022b). Only 7 % of studies have investigated this approach under greenhouse conditions (Figure 3), underscoring the need for further research in this sense.
On the other hand, bioactive compounds in botanical extracts tend to oxidize or degrade over time, limiting the duration of their beneficial effects on plants (Ali et al., 2021). Therefore, applications should be every 7 to 14 days to obtain an optimal response in crops (Brazales-Cevallos et al., 2022; Velasco-Ramírez et al., 2022a; Garza-García et al., 2023; Jasso de Rodríguez et al., 2024a; Moreno-Guerrero et al., 2024). In addition, the efficacy of the extracts also depends on the de- velopmental stage of the crop, with transplanting, flowering, and fruiting being key moments for their application (Jasso de Rodríguez et al., 2020; 2023).
The analyzed studies used 12 crop species as experimental models to evaluate the biostimulant properties of botanical extracts. Among these, horticultural crops were the most extensively studied, accounting for 67 % of the research included in this review. Cucumber (Cucumis sativus L.) was the most frequently studied crop, appearing in 27 % of the publications. Other key vegetables, such as tomato (Solanum lycopersicum L.) and lettuce (Lactuca sativa L.), were evalua- ted in 20 % of the studies. Bell pepper (Capsicum annuum L.) was addressed in 13 % of the publications, while carrot (Daucus carota L.) and radish (Raphanus sativus L.) were each featured in only 7 %.
This preference for leaf, root, or fruit vegetables is pro- bably attributed to their short growth cycle, which facilitates the observation of rapid treatment responses. This group of crops has been recommended in NOM-077-FITO-2000 to carry out biological effectiveness studies of plant nutrition inputs (DOF, 2020). In addition, vegetables represent a fun- damental part of agricultural production in Mexico, where approximately 20.3 % of arable areas are used for this type of crop (SIAP, 2024).
On the other hand, 7 % of the studies investigated the biostimulant effects of botanical extracts on fruit (Fragaria ananassa Duch.), ornamental (Catharanthus roseus L.), and medicinal (Calendula officinalis L.) crops. The remaining 20
% of the published articles included crops such as Salicornia bigelovii Torr., Stevia rebaudiana Bertoni, and Arabidopsis thaliana L., species with various agroindustrial and biological applications.
Several studies in recent years have demonstrated the posi- tive effects of crude botanical extracts or their organic frac-
tions on seed germination, vegetative growth, flowering, and fruiting of crops of commercial interest (Jasso de Rodríguez et al., 2020; Velasco-Ramírez et al., 2021; Jasso de Rodríguez et al., 2023; Velasco-Ramírez et al., 2022b; Alcalán-López et al., 2023; Jasso de Rodríguez et al., 2024a; Moreno-Guerrero et al., 2024; Velázquez-Gamboa et al., 2024; Godlewska et al., 2021; Han et al., 2024). These processes are critical to agricul- tural production, directly impacting crop yield and quality (Martínez-Lorente et al., 2024). Table 2 summarizes the biosti- mulant effects of Mexican plant extracts and the metabolites responsible for their biological activities.
Seed germination is a key stage for agricultural production because it marks the beginning of plant growth and largely determines crop yield (Carvalho et al., 2022). Therefore, it is important to implement natural strategies that ensure suc- cessful and uniform germination to obtain vigorous plants resistant to various environmental conditions (Gupta et al., 2022). Seed treatment (i.e., imbibition or soaking) with bota- nical extracts is a simple and effective technique to improve
germination and stimulate seedling growth (Martínez-Lo- rente et al., 2024). However, the concentration of the extracts evaluated is a determining factor in their effectiveness as biostimulants (Han et al., 2024).
For example, liquid extracts obtained from different halophytes at 5 % increased the relative germination percentage, relative radicle growth, seedling length, and germination index of crops such as carrot (D. carota), lettuce (L. sativa), radish (R. sativus), and cucumber (C. sativus). High concentrations (20 %) significantly reduced these parame- ters (Alcalán-López et al., 2023). In another study, aqueous extracts of V. sphaerocephala and V. fastigiata at 0.5 % and
1.0 % adversely affected the germination percentage, mean germination time, and vigor index of tomato and cucumber seeds. However, they promoted the growth of leaves and seedling roots of these potted vegetables (Velasco-Ramírez et al., 2023). These authors suggested that Verbesina extracts contain allelopathic compounds that inhibit germination. On the other hand, Jasso de Rodríguez et al. (2024b) reported the highest values of total plumule and radicle length, as well as the fresh weight of seedlings of two tomato varieties, at
Table 2. Biostimulant effects of extracts obtained from plants distributed in Mexico, and responsible metabolites identified in the study * or suggested from the literature **.
Tabla 2. Efectos bioestimulantes de extractos obtenidos de plantas distribuidas en México, y metabolitos responsables identificados en el estudio * o sugeridos en la literatura **.
Plant source for extract | Responsible metabolites | Biostimulant effect | Plant evaluated | References |
Grasses | Humic substances** | Improved seed germination and radicle growth under salt stress | Chili pepper | (Huez-López et al., 2008) |
Myriophyllum aquaticum | Phenolic compounds* | Increased root elongation | Lettuce | (Viveros-Legorreta et al., 2018). |
Rhus trilobata, Rhus muelleri, Flourensia microphylla, Flourensia retinophylla, Cucurbita foetidissima | Phenolic compounds** | Increased vegetative growth, crop yield, and fruit quality | Tomato | (Jasso de Rodríguez et al., 2020) |
Verbesina sphaerocephala | Phenolic compounds* | Stimulated vegetative growth and flowering | Cucumber | (Velasco-Ramírez et al., 2021) |
Moringa oleifera | Flavonoids, Amino Acids** | Induce tolerance to salinity | Arabidopsis | (Brazales-Cevallos et al., 2022) |
Verbesina sphaerocephala Verbesina fastigiata | Phenolic compounds** | Promoted seedling growth | Tomato, cucum- ber | (Velasco-Ramírez et al., 2022a) |
Verbesina sphaerocephala, Verbesina fastigiata | Phenolic compounds** | Increased vegetative growth, flowering, and fruit production | Strawberry | (Velasco-Ramírez et al., 2022b) |
Bacopa monieri, Distichlis spicata, Heliotropium curasavicun, Flaveria trinervia, Sesuvium verrucosum | Not mentioned | Increased germination para- meters | Carrot, lettuce, radish, and cu- cumber | (Alcalán López et al., 2023) |
Maclura tinctoria | Phenolic compounds* | Stimulation of plant growth, height, and flowering | Salicornia bigelovii | (Corona et al., 2023) |
Amphipterygium glaucum | Flavonoids, polyphenols, reducing sugars* | Improved vegetative develop- ment, flower production, and photosynthetic pigment content | Vinca, Calendula | (Garza-García et al., 2023) |
R. muelleri, Rhus virens, F. microphylla, F. retinophylla | Phenolic compounds** | Promotion of growth, yield, and fruit quality | Bell pepper | (Jasso de Rodríguez et al., 2023) |
R. muelleri, F. microphylla F. retino- phylla | Phenolic compounds** | Increased growth, yield, and fruit quality | Cucumber | (Jasso de Rodríguez et al., 2024a) |
R. muelleri, R. virens, Jatropha dioica, Juglans mollis | Phenolic compounds** | Stimulated root, stem, and leaf growth | Tomato | (Jasso de Rodríguez et al., 2024b) |
Opuntia ficus-indica, Coriandrum sativum | Phenolic compounds, phytohormones** | Increased mineral nutrient content | Lettuce | (Moreno-Guerrero et al., 2024) |
Moringa oleifera | Not mentioned | Increased phenolic compounds and glycosides content in leaves | Stevia rebaudiana | (Velazquez-Gamboa et al., 2024) |
the lowest (75 mg mL–1) and highest (200 mg mL–1) concen- trations of semi-desert plant extracts.
Also, the application of botanical extracts has been shown to improve the germination percentage and growth of bell pepper (C. annuum) and Arabidopsis thaliana seed- lings under salinity stress conditions (Huez-Lopez et al., 2008; Brazales-Cevallos et al., 2022). In general, the variability in ob- served effects across studies can be attributed to differences in extract type, the plant species or cultivar evaluated, and the experimental conditions (Carvalho et al., 2022).
Enhancing crop morphology and physiology during the growth phase is associated with increased productivity, which is the primary objective of agricultural production (Zulfiqar et al., 2020). Appropriate treatment with botanical extracts at different phenological stages of crops has positive effects on growth and productivity parameters (Godlewska et al., 2021; Han et al., 2024).
In a study, extracts of R. muelleri and J. dioica stimulated root and aerial growth of tomato seedlings under green- house conditions, more than commercial rooting products (Jasso de Rodríguez et al., 2024b). Similarly, another study reported that the application of botanical extracts increa- sed stem length and diameter, as well as leaf dry weight, in tomato plants. Moreover, the extract of R. muelleri showed greater efficacy in increasing the number, weight, and yield of tomatoes, reaching values similar to those obtained with the bioregulator 6-benzylaminopurine (Jasso de Rodríguez et al., 2020). Likewise, applying F. retinophylla extract to bell pepper plants has stimulated longitudinal stem growth and enhanced total dry matter accumulation and root volume. This, in turn, increased fresh weight, volume, and yield of pepper fruit (Jasso de Rodríguez et al., 2023).
The biostimulant effect of botanical extracts, whether applied individually or in combination, has also been de- monstrated to improve growth parameters (e.g., plant height, leaf number, chlorophyll index, and biomass accumulation) and enhance the productivity of cucumber plants (Velasco- Ramírez et al., 2021; Jasso de Rodríguez et al., 2024a).
A study by Velasco-Ramírez et al. (2022b) found that extracts of V. sphaerocephala and V. fastigiata improved the growth (leaf width and length, crown number, and root length) and reproductive development (number of buds, flowers, and fruits) of strawberry plants grown in pots. On the other hand, the foliar application of nanoparticles synthe- sized from the methanolic extract of A. glaucum stimulated better vegetative development and increased the content of photosynthetic pigments in vinca (C. roseus) and calendula (C. officinalis) plants (Garza-García et al., 2023). In contrast, nopal (O. ficus-indica) and coriander (C. sativum) extracts did not positively affect the growth of lettuce plants (Moreno- Guerrero et al., 2024).
In recent years, there has been a notable increase in the
prevalence of abiotic stress conditions in agriculture, largely due to the effects of climate change (Ali et al., 2021; Martínez- Lorente et al., 2024). These situations induce physiological stress that can significantly reduce crop yields (Ahmad et al., 2022). The use of plant-derived biostimulants is emerging as a cost-effective and sustainable strategy to mitigate the effects of adverse environmental conditions on crops (Godlewska et al., 2021; Ahmad et al., 2022; Martínez-Lorente et al., 2024). However, research on the potential of Mexican plant extracts to enhance abiotic stress tolerance is still in its early stages.
The study by Huez-Lopez et al. (2008) reported that C. echinatus extract, obtained by fermentation, improved seed germination percentage and the length of the radicle and hy- pocotyl of chili pepper (C. annuum) seedlings under salinity conditions. Likewise, Brazales-Cevallos et al. (2022) observed that M. oleifera leaf extracts induce tolerance to saline stress on A. thaliana seeds. This study offers significant insights into the mechanisms at the transcriptomic level underlying the protective effects of plant-based biostimulants during the initial stages of plant development.
Ensuring access to sufficient, safe, and high-quality nutritious food to meet dietary needs represents one of the major challenges facing agriculture in the quest for food security. The quality of agricultural products is a key factor influencing their commercial value, consumer satisfaction, and the reduc- tion of post-harvest losses (Akter et al., 2024). The concept of quality encompasses the physical appearance of the product (i.e., color, size, shape, firmness) as well as the organoleptic, nutritional and nutraceutical properties that determine its acceptability and health benefits (Reis et al., 2022). A recently recognized strategy for enhancing crop quality is the applica- tion of plant biostimulants (Calvo et al., 2014; Rouphael and Colla, 2020; Han et al., 2024). In this regard, several studies included in this review highlight the potential of botanical extracts to improve various quality attributes, demonstrating their capacity to positively influence the characteristics of the final product.
The foliar application of extracts from plants native to the semiarid regions of northeastern Mexico resulted in tomato fruits (S. lycopersicum) with polar length, equatorial diameter, firmness, pH, total soluble solids (TSS), lycopene, and vitamin C content comparable to those obtained with the synthetic cytokinin 6-BAP (Jasso de Rodríguez et al., 2020). Similarly, fo- liar spraying of semi-desert plant extracts enhanced several quality attributes of bell peppers, including longitudinal and equatorial diameters, volume, TSS, anthocyanin, and vitamin C content. The values obtained were comparable to or excee- ded, those obtained with commercial bioregulators (Jasso de Rodríguez et al., 2023). In another study, combining various botanical extracts improved several of the aforementioned quality parameters in cucumber fruits (Jasso de Rodríguez et al., 2024a). On the other hand, S. rebaudiana plants treated with nanoparticles synthesized from the aqueous extract of
M. oleifera showed a higher leaf content of phenolic com-
pounds (chlorogenic acid and caffeic acid) and glycosides (Velázquez-Gamboa et al., 2024). Moreover, foliar application of nopal and coriander extracts increased the concentra- tion of macro and micronutrients in the lettuce leaf tissue (Moreno-Guerrero et al., 2024).
According to Yakhin et al. (2017), the mode of action refers to the “specific effect on a discrete biochemical or regulatory process”, while the mechanism of action involves the “impact on general biochemical or molecular pathways or physiologi- cal processes”. The inherent chemical complexity of botanical extracts has thus far precluded the requisite level of characte- rization necessary to elucidate their specific modes of action. Therefore, their functions in plants are mainly inferred from the underlying mechanisms of action (Martínez-Lorente et al., 2024).
In this regard, the overall positive effects of the bota- nical extracts on plant growth and productivity are due to the modulation of various physiological and biochemical processes, such as (1) nutrient and water uptake, (2) pho- tosynthetic activity, (3) modulation of phytohormones, (4) specialized and oxidative metabolism, (5) gene expression related to abiotic stress tolerance, and (6) plant architecture and phenology (Zulfiqar et al., 2020; Godlewska et al., 2021; Ahmad et al., 2022; Martínez-Lorente et al., 2024). However, it should be clarified that the biostimulatory properties of botanical extracts on plant performance are not the result of a single bioactive compound, but of the coordinated and synergistic action of predominant compounds, such as po- lyphenols, phytohormones, amino acids, sterols, terpenoids, and vitamins (Ahmad et al., 2022; Han et al., 2024; Jasso de Rodríguez et al., 2024a). Although some studies have repor- ted the presence of micro- and macronutrients in plants- derived extracts (Huez-Lopez et al., 2008; Velasco-Ramírez et al., 2022b; Alcalán-López et al., 2023), their concentrations are often insufficient to induce physiological responses in plants at the doses commonly employed (Han et al., 2024). This observation suggests that organic constituents, rather than mineral elements, are primarily responsible for growth- stimulating activity.
Most of the studies analyzed in this review attribute the beneficial effects of extracts from Mexican plants to their phe- nolic compound content. These compounds represent one of the most abundant families of secondary plant metabolites and include phenolic acids, flavonoids, tannins, and others (Michalak et al., 2006; Martínez-Lorente et al., 2024). Recently, interest in phenolics has increased considerably due to their potential as biostimulants in agriculture (Rouphael and Colla, 2020). Phenolics act as potent antioxidants because they can scavenge reactive oxygen species, thus helping to maintain redox homeostasis and protect cells from oxidative damage under normal and stressful conditions (Michalak et al., 2006). Furthermore, they modulate the metabolism and transport of auxins and ethylene, thereby indirectly influencing the
processes regulated by these phytohormones (Santner et al., 2009). For example, auxins are primarily associated with growth and development, while ethylene regulates matu- ration, senescence, and stress response (Ali et al., 2021). By the above, several studies on plant methanolic extracts in various horticultural crops have reported biostimulatory activity comparable to that of synthetic bioregulators, such as indoleacetic acid, gibberellic acid, and 6-benzylaminopu- rine (Jasso de Rodríguez et al., 2020; Jasso de Rodríguez et al., 2024a; Jasso de Rodríguez et al., 2024b). In another study, moringa leaf extracts induced tolerance to salinity stress in A. thaliana seeds through the transcriptional activation of abs- cisic acid, salicylic acid, auxin, and ethylene-related signaling pathways (Brazales-Cevallos et al., 2022).
Additionally, phenolic compounds can stimulate pho-
tosynthesis, enhance CO2 fixation, and regulate the stomatal opening/closing mechanism, promoting better plant adap- tation to environmental conditions (Ertani et al., 2016; Agati et al., 2012; Martínez-Alonso et al., 2022). On the other hand, when applied to the soil, flavonoids can function as chelating agents for metal ions, enhancing their availability for plant nutrition (Cesco et al., 2012). Moreover, it has been propo- sed that flavonoids released into the rhizosphere stimulate the solubilization of inorganic phosphorus from the soil, allowing its plant uptake (Tomasi et al., 2008). These phenolic compounds have also been found to increase the expression of aquaporins in biological membranes, improving water uptake efficiency (Martinez-Alonso et al., 2022).
Another important role of phenolics is their participa- tion in plant-microorganism interactions (Badri et al., 2009). It has been described that nod genes, which are activated in legumes during symbiosis with nitrogen-fixing bacteria, are stimulated by flavonoids acting as chemical signals that tri- gger their activation (Cooper, 2007). Furthermore, both flavo- noids and some phenolic acids have been shown to interfere with quorum sensing, a microbial communication system used in symbiosis, defense, and other plant-microorganism interactions (Zhuang et al., 2005).
Finally, based on the scientific reports included in this work, we suggest that the bioactive compounds in botani- cal extracts can promote plant growth and increase plant productivity (yield and quality) through direct and indirect mechanisms of action. When absorbed through the leaves or roots, some compounds interact directly with plant me- tabolism and physiology. In contrast, others act indirectly by improving soil properties, enhancing the rhizospheric environment, and stimulating beneficial soil microbiota.
Biostimulants derived from native plant extracts are currently experiencing a global surge in interest, aiming to valorize local natural resources to enhance agriculture and promote food sovereignty (Godlewska et al., 2021; Han et al., 2024).
However, the studies analyzed in this review show that only a limited number of plant species in Mexico have been eva- luated for this purpose. In this context, the systematic search for natural compounds and whole organisms with potential commercial value—known as bioprospecting—is a valuable tool for increasing our understanding of species with known biostimulant properties (Torres and Velho, 2009). In the con- text of this publication, a bioprospecting was focused on the identification and development of novel plant-derived bio- active compounds for use as agricultural biostimulants, and a methodology in two-phases was proposed. To achieve this, the integration of knowledge, technologies and processes is needed through schemes with systematic approaches and phases such as the one presented in Figure 4.
Phase 1 focuses on the initial exploration and selection of species of interest with agricultural potential. The process starts with a judicious assessment of potential plant species, weighing their favourable attributes (such as abundance in the ecosystem, ease of access and their ability to thrive in challenging environmental conditions) against potential constraints, such as biomass yield or the stability of their compounds. Once a species has been pre-selected, the re- search is aimed at locating the metabolites of interest within the plant, commonly concentrated in the leaves. Priority is given to aqueous extraction methods due to their simplicity and applicability both in the laboratory and in the field, also
incorporating a sustainability perspective when considering the use of residual biomass. Preliminary biological assays, such as germination tests or rapid growth models, are then performed to evaluate the biostimulant potential of the extracts obtained. This stage requires previously defining the key variables that will allow the effect to be measured, such as germination growth, and vigor parameters of the plants. It is important that bioprospecting studies consider the specifi- cations of NOM-077-FITO-2000, which are mainly applicable in phase 1 of this strategy (Figure 4). Once the exploratory phase is over, Phase 2 delves into scientific research and product development. It begins with an exhaustive chemical characterization of those extracts that showed promising activity, in order to identify the molecules responsible for the biostimulant effect. Accurate quantification of these active molecules, both in the extracts and in the original plant, is another crucial step in standardizing and understanding the dose-response. At the same time, it seeks to elucidate the underlying mechanisms of action, that is, how these molecules interact at a biochemical, physiological or genetic level in the plant. Subsequently, the research addresses the ways for the formulation of the product, will delve into the methods used to create or develop a product. At this point the research will explore the specific techniques, processes, and approaches involved in taking an idea and turning it into a tangible product up to sustainable and scalable production
Figure 4. Phases of bioprospecting of plant species for the production biostimulants.
Figura 4. Fases de bioprospección de especies vegetales para la producción de bioestimulantes.
of the bioinput, exploring options such as the cultivation of the source species, the culture of plant tissues or even the organic synthesis of the identified molecules. Prior to final implementation, a rigorous cost-benefit analysis is perfor- med to determine the economic viability of the technology. Finally, the process of registering intellectual property is contemplated to protect the innovation developed.
Interest in plant biostimulants has increased significantly in recent years, driven by the need to adopt more sustainable and efficient agricultural practices. However, there are still se- veral issues to be addressed, mainly related to their definition and regulation. In Mexico, the production, commercializa- tion, and use of biostimulants in agriculture remain limited, partly due to the absence of a specific regulatory framework. Currently, these products attempt to be regulated indirectly through the Norma Oficial Mexicana NOM-077-FITO-2000, which focuses on plant nutrition inputs, such as organic soil improvers, inoculants, and type 1 and 2 growth regu- lators. Therefore, a coordinated effort between academia, government regulatory agencies, and industry is needed to harmonize current regulations and enhance farmers’ com- prehension of the functions and benefits of biostimulants.
Among the diversity of raw materials recognized for biostimulant production, plants stand out for their accessibi- lity, wide availability, and richness in primary and secondary metabolites with multiple biological functions. In this sense, the studies analyzed in this review support the biostimulant potential of extracts from plant species distributed in Mexi- co. However, despite the remarkable floristic richness of the country, few species have been studied for this purpose. In particular, the Asteraceae family concentrates the largest number of species evaluated in bioprospecting studies.
The application of botanical extracts to seeds, plants, or soil has been demonstrated to have positive effects on key processes such as germination, seedling vigor, vegetative growth, reproductive development, abiotic stress tolerance, as well as on crop yield and nutritional quality. Horticultural crops are among the most used experimental models in studies evaluating the biological effectiveness of botanical extracts, both under laboratory and greenhouse conditions. Most studies attribute the observed benefits to phenolic compounds, including phenolic acids and flavonoids. However, the efficacy of these extracts can be influenced by several factors, including the characteristics of the raw material (species, phenological stage, part of the plant), the extraction method (extraction technique and solvent), and the conditions of application to the plant (concentration, dose, frequency, and timing of application).
Although advances in the understanding of Mexican plant extracts highlight their potential as a promising stra- tegy for sustainable crop management aimed at enhancing production and productivity, further research is still required. The exploration of new species with biostimulant properties, especially those native or endemic species with the ability
to adapt to specific environmental conditions, would allow the identification of novel biological activities and bioactive compounds. Moreover, research on the mechanism of action requires the elucidation of the chemical compounds respon- sible for the effects observed in botanical extracts already studied, as well as a further understanding of the physiologi- cal, metabolic, and genomic modifications induced in plants. Likewise, to optimize the use of these bioproducts in agricul- ture, crop-specific research is needed to study in depth the duration of the effect after the application of the extract, the most appropriate phenological stages for its application and maximization of benefits, as well as the possibility of obtai- ning synergistic effects by combining extracts from different plant species or concentrations. Additional trials also should be conducted under field conditions to demonstrate the efficacy of these botanical formulations in natural settings, including abiotic stress situations. Finally, using advanced biotechnological tools, such as nanotechnology, can enhance the biostimulant properties of botanical extracts, improving their stability, bioavailability, and competitiveness against synthetic agro-inputs.
The authors declare no conflict of interest.
Agati, G., Azzarello, E., Pollastri, S. and Tattini, M. 2012. Flavonoids as antioxidants in plants: Location and functional significance. Plant Science. 196: 67–76. https:// doi.org/10.1016/J.PLANTSCI.2012.07.014
Ahmad, A., Blasco, B. and Martos, V. 2022. Combating salinity through natural plant extracts based biostimulants: A review. Frontiers in Plant Science. 13: 862034. https://doi. org/10.3389/fpls.2022.862034
Akter, T., Bhattacharya, T., Kim, J.H., Kim, M.S., Baek, I., Chan, D.E. and Cho, B.K. 2024. A comprehensive review of external quality measurements of fruits and vegetables using nondestructive sensing technologies. Journal of Agriculture and Food Research. 15: 101068. https://doi.org/10.1016/J. JAFR.2024.101068
Ali, O., Ramsubhag, A. and Jayaraman, J. 2021. Biostimulant properties of seaweed extracts in plants: Implications towards sustainable crop production. Plants. 10(531): 1–27. https://doi.org/10.3390/plants10030531
Alcalán-López, L.Á., Lastiri-Hernández, M.A. and Álvarez-Bernal,
D. 2023. Efecto de biofertilizantes líquidos, obtenidos a partir de halófitas, en la germinación y emergencia de cuatro especies de hortalizas (Daucus carota L., Lactuca sativa L., Raphanus sativus L. y Cucumis sativus L.). Biotecnia. 25(3): 197–207. https://doi.org/10.18633/biotecnia.v25i3.2127
Badri, D.V., Weir, T.L., van der Lelie, D. and Vivanco, J. M. 2009. Rhizosphere chemical dialogues: plant–microbe interactions. Current Opinion in Biotechnology. 20(6): 642–650. https://doi.org/10.1016/J.COPBIO.2009.09.014
Banco de México [accessed February 1, 2025]. 2022. Estimación de la exposición regional ante cambios en el mercado de fertilizantes nitrogenados. Available online at: https://www. banxico.org.mx/publicaciones-y-prensa/reportes-sobre- las-economias-regionales/recuadros/%7B3F3D3D32-A106- FC6B-5746-0BC00AA60666%7D.pdf
Bonilla-Barbosa, J.R. and Santamaría, B. 2013. Plantas acuáticas exóticas y traslocadas invasoras. En Especies acuáticas invasoras en México. R. Mendoza y P. Koleff (ed.), pp. 223–247. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México.
Brazales-Cevallos, D.K., Romero-Contreras, Y.J., Vences-Guzmán, M.Á., Torres, M., Aviles-Baltazar, N.Y., Sohlenkamp, C., and Serrano, M. 2022. Transcriptional characterization of the biostimulant effect of Moringa oleifera leaf extracts using Arabidopsis thaliana as a model. South African Journal of Botany, 144, 250–256. https://doi.org/10.1016/j. sajb.2021.09.011
Calvo, P., Nelson, L. and Kloepper, J.W. 2014. Agricultural uses of plant biostimulants. Plant and Soil. 383: 3–41. https://doi. org/10.1007/s11104-014-2131-8
Caradonia, F., Battaglia, V., Righi, L., Pascali, G. and La Torre, A. 2019. Plant biostimulant regulatory framework: Prospects in Europe and current situation at international level. Journal of Plant Growth Regulation. 38: 438–448. https://doi. org/10.1007/s00344-018-9853-4
Carvalho, R. da S., Silva, M.A. da, Borges, M.T.M.R. and Forti, V.A. 2021. Plant extracts in agriculture and their applications in the treatment of seeds. Ciência Rural, 52(5), e20210245. https://doi.org/10.1590/0103-8478cr20210245
Cesco, S., Mimmo, T., Tonon, G., Tomasi, N., Pinton, R., Terzano, R., Neumann, G., Weisskopf, L., Renella, G., Landi, L. and Nannipieri, P. 2012. Plant-borne flavonoids released into the rhizosphere: Impact on soil bioactivities related to plant nutrition. A review. Biology and Fertility of Soils. 48(2): 123–149. https://doi.org/10.1007/s00374-011-0653-2
Cooper, J.E. 2007. Early interactions between legumes and rhizobia: disclosing complexity in a molecular dialogue. Journal of Applied Microbiology. 103(1): 1355–1365.
Corona, B.E.L., Ocampo, A.G., Juárez, D.R., García, J.O., Fernández,
I.M. and Puente, E.O.R. 2023. Biostimulant effect of chitosan and phenolic extracts on the phenological development of the halophyte Salicornia bigelovii (Torr.). Journal of the Saudi Society of Agricultural Sciences. 22(8): 584–590. https://doi. org/10.1016/j.jssas.2023.08.001
Corsi, S., Ruggeri, G., Zamboni, A., Bhakti, P., Espen, L., Ferrante, A., Noseda, M., Varanini, Z. and Scarafoni, A. 2022. A bibliometric analysis of the scientific literature on biostimulants. Agronomy. 12(6): 1257. https://doi. org/10.3390/agronomy12061257
DataMéxico [accessed February 1, 2025]. 2025. Fabricación de fertilizantes, pesticidas y otros agroquímicos. Available online at: https://www.economia.gob.mx/datamexico/es/ profile/industry/pesticide-fertilizer-and-other-agricultural- chemical-manufacturing
DOF, Diario Oficial de la Federación. [accessed: January 30, 2025]. 2020. Norma Oficial Mexicana NOM-077-FITO-2000. Estudios de efectividad biológica en insumos de nutrición vegetal de uso agrícola y su Dictamen Técnico. Available online at: https://www.dof.gob.mx/nota_detalle.php?codig o=5592222&fecha=24/04/2020#gsc.tab=0
DOF, Diario Oficial de la Federación. [accessed January 30, 2025]. 2010. NORMA Oficial Mexicana NOM-059-SEMARNAT-2010,
Protección ambiental-Especies nativas de México de flora y fauna silvestres-Categorías de riesgo y especificaciones para su inclusión, exclusión o cambio-Lista de especies en riesgo. Available online at: https://www.dof.gob.mx/ normasOficiales/4254/semarnat/semarnat.htm
Du Jardin, P. 2015. Plant biostimulants: Definition, concept, main categories and regulation. Scientia Horticulturae. 196: 3–14. https://doi.org/10.1016/j.scienta.2015.09.021
Ertani, A., Pizzeghello, D., Francioso, O., Tinti, A. and Nardi, S. 2016. Biological activity of vegetal extracts containing phenols on plant metabolism. Molecules. 21(2): 205. https:// doi.org/10.3390/MOLECULES21020205
European Union. [accessed December 17, 2024]. 2019. Regulation (EU) 2019/1009 of the European Parliament and of the Council of 5 June 2019 laying down rules on the placing of EU fertilising products on the market and amending Regulations (EC) No 1069/2009 and (EC) No 1107/2009 and repealing Regulation (EC) No 2003/2003. Available at: https://eur-lex.europa.eu/legal-content/EN/ TXT/?uri=CELEX:32019R1009
Garza-García, J.J.O., Hernández-Díaz, J.A., León-Morales, J.M., Velázquez-Juárez, G., Zamudio-Ojeda, A., Arratia-Quijada, J., Reyes-Maldonado, O.K., López-Velázquez, J.C. and García-Morales, S. 2023. Selenium nanoparticles based on Amphipterygium glaucum extract with antibacterial, antioxidant, and plant biostimulant properties. Journal of Nanobiotechnology. 21: 252. https://doi.org/10.1186/ s12951-023-02027-6
Godlewska, K., Ronga, D. and Michalak, I. 2021. Plant extracts- importance in sustainable agriculture. Italian Journal of Agronomy. 16(2): 1851. https://doi.org/10.4081/ ija.2021.1851
GPO, United States Government Publishing Office. [accessed: December 20, 2024] 2023. To amend the Federal Insecticide, Fungicide, and Rodenticide Act to provide for a consistent definition for plant biostimulants. Available at: https://www. govinfo.gov/app/details/BILLS-118s802is
Gupta, S., Doležal, K., Kulkarni, M.G., Balázs, E. and Van Staden,
J. 2022 Role of non-microbial biostimulants in regulation of seed germination and seedling establishment. Journal of Plant Growth Regulation. 97: 271–313. https://doi. org/10.1007/s10725-021-00794-6
Han, M., Kasim, S., Yang, Z., Deng, X., Saidi, N.B., Uddin, M.K. and Shuib, E.M. 2024. Plant extracts as biostimulant agents: A promising strategy for managing environmental stress in sustainable agriculture. Phyton-International Journal of Experimental Botany. 93(9): 2149–2166. https://doi. org/10.32604/phyton.2024.054009
Huez-Lopez, M.A., Samani, Z., Elías, J.L., Avilés, A.Á. and Flores,
F.P. 2008. Efecto de un extracto vegetal en la germinación de semillas de chile (Capsicum annuum L.) bajo condiciones salinas. Biotecnia. 10(3): 11–19.
INEGI, Instituto Nacional de Estadística y Geografía. [accessed February 1, 2025] 2019. Encuesta Nacional Agropecuaria 2019. Available at: https://www.inegi.org.mx/contenidos/ programas/ena/2019/doc/rrdp_ena2019.pdf
Jasso de Rodríguez, D., Alonso-Cueva, C.F., Rodríguez-García, R., Ramírez, H., Díaz-Jiménez, L., Villareal-Quintanilla, J.A. and Juárez-Maldonado, A. 2020. Extractos de plantas del semidesierto en la inducción del crecimiento de tomate (Lycopersicon esculentum Mill). Ecosistemas y Recursos Agropecuarios. 7(1): e2342. https://doi.org/10.19136/era. a7n1.2342
Jasso de Rodríguez, D., Rocha-Rivera, M.F., Ramírez-Rodríguez, H., Villarreal-Quintanilla, J.Á., Díaz-Jiménez, M.L.V., Rodríguez- García, R. and Carrillo-Lomelí, D.A. 2023. Extractos de plantas
como bioestimulantes de crecimiento, rendimiento y calidad de fruto en pimiento morrón. Ecosistemas y Recursos Agropecuarios. 10(2): e3559. https://doi.org/10.19136/era. a10n2.3559
Jasso de Rodríguez, D., Cuevas-Julio, I., Ramírez-Rodríguez, H., Villarreal Quintanilla, J.Á., Hernández-Pérez, A., Díaz- Giménez, M.L.V., Peña Ramos, F.M. and Carrillo Lomelí,
D. A. 2024a. Bioestimulaciónde extractos de plantas del semidesierto en crecimiento, rendimiento y calidad en pepino. Ecosistemas y Recursos Agropecuarios. 11(3): e4186. https://doi.org/10.19136/era.a11n3.4186
Jasso de Rodríguez, D., Ramírez-Pérez, C., Ramírez-Rodríguez, H., Villarreal-Quintanilla, J.Á., Hernández-Pérez, A., Díaz- Jimenez, M.L.V. and Peña-Ramos, F.M. 2024b. Bioestimulantes de plantas del semidesierto en el crecimiento radicular y aéreo de plántulas de tomate. Ecosistemas y Recursos Agropecuarios. 11(3): e4145. https://doi.org/10.19136/era. a11n3.4145
Leopardi-Verde, C.L., Guzmán-González, S., Carnevali, G., Duno de Stefano, R. and Tapia-Muñoz, J.L. 2021. Weeds of commercial crops in Colima, Mexico. Revista Mexicana de Biodiversidad. 92: e923622. https://doi.org/10.22201/ ib.20078706e.2021.92.3622
Martinez-Alonso, A., Garcia-Ibañez, P., Bárzana, G. and Carvajal,
M. 2022. Leaf gas exchange and growth responses of tomato plants to external flavonoids application as biostimulators under normal and salt-stressed conditions. Agronomy. 12(12): 3230. https://doi.org/10.3390/AGRONOMY12123230
Martínez-Lorente, S.E., Martí-Guillén, J.M., Pedreño, M.Á., Almagro, L. and Sabater-Jara, A.B. 2024. Higher plant- derived biostimulants: Mechanisms of action and their role in mitigating plant abiotic stress. Antioxidants. 13: 318. https://doi.org/10.3390/antiox13030318
Mattner, S.W., Wite, D., Riches, D.A., Porter, I.J. and Arioli, T. 2013. The effect of kelp extract on seedling establishment of broccoli on contrasting soil types in southern Victoria, Australia. Biological Agriculture & Horticulture. 29(4): 258–270. https://doi.org/10.1080/01448765.2013.830276
Michalak, A. 2006. Phenolic compounds and their antioxidant activity in plants growing under heavy metal stress. Polish Journal of Environmental Studies. 15(4): 523–530.
Michalak, I. and Chojnacka, K. 2014. Algae extracts: Technology and advances. Engineering in Life Sciences. 14(6): 581–591. https://doi.org/10.1002/elsc.201400139
Moreno-Guerrero, D.E., Ramírez-Olvera, S.M., Ojeda-Salgado, H.L., Pérez-Mercado C.A. and Trejo-Téllez, L.I. 2024. Effect of plant extracts in growth and nutrimental concentration of lettuce leaves (Lactuca sativa). Biotecnia. 26: e1929. https:// doi.org/10.18633/biotecnia.v26.1929.
Pérez-Alva, A., MacIntosh, A.J., Baigts-Allende, D.K., García-Torres,
R. and Ramírez-Rodrigues, M.M. 2022. Fermentation of algae to enhance their bioactive activity: A review. Algal Research. 64: 102684. https://doi.org/10.1016/j.algal.2022.102684
Ricci, M., Tilbury, L., Daridon, B. and Sukalac, K. 2019. General principles to justify plant biostimulant claims. Frontiers in Plant Science. 10: 494. https://doi.org/10.3389/ fpls.2019.00494
Reis, F.R., Marques, C., de Moraes, A.C.S. and Masson, M.L. 2022. Trends in quality assessment and drying methods used for fruits and vegetables. Food Control. 142: 109254. https:// doi.org/10.1016/J.FOODCONT.2022.109254
Rouphael, Y. and Colla, G. 2020. Editorial: Biostimulants in agriculture. Frontiers in Plant Science. 11: 40. https://doi. org/10.3389/fpls.2020.00040
Sanjuán, J., Nápoles, M.C., Pérez-Mendoza, D., Lorite, M.J. and Rodríguez-Navarro, D.N. 2023. Microbials for agriculture: Why do they call them biostimulants when they mean probiotics?. Microorganisms. 11(1): 153. https://doi. org/10.3390/microorganisms11010153
Santner, A., Calderon-Villalobos, L. and Estelle, M. 2009. Plant hormones are versatile chemical regulators of plant growth. Nature Chemical Biology. 5: 301–307. https://doi. org/10.1038/nchembio.165
SEMARNAT, Secretaría de Medio Ambiente y de Recursos Naturales. [accessed February 2, 2025]. 2012. Informe de la situación del medio ambiente en México. Available at: https://apps1.semarnat.gob.mx:8443/dgeia/informe_ 12/04_biodiversidad/cap4_1.html
SIAP, Servicio de Información Agroalimentaria y Pesquera. [accessed December 18, 2024]. 2024. Cierre de la producción agrícola. Anuario Estadístico de la Producción Agrícola. México. Servicio de Información Agrícola y Pesquera. Available at: https://nube.siap.gob.mx/cierreagricola/
Tomasi, N., Weisskopf, L., Renella, G., Landi, L., Pinton, R., Varanini, Z. and Cesco, S. 2008. Flavonoids of white lupin roots participate in phosphorus mobilization from soil. Soil Biology and Biochemistry. 40(7): 1971–1974. https://doi. org/10.1016/j.soilbio.2008.02.017
Torres, O.D. and Velho, L. 2009. La bioprospección como un mecanismo de cooperación internacional para fortalecimiento de capacidades en ciencia y tecnología en Colombia. Ciência da Informação 38(3): 96–110. https://doi. org/10.1590/s0100-19652009000300007
Velasco-Ramírez, A.P., Velasco-Ramírez, A., Hernández-Herrera, R.M., Ceja-Esquivez, J., Velasco-Ramírez, S.F., Ramírez- Anguiano, A.C. and Torres-Morán, M.I. 2022a. The impact of aqueous extracts of Verbesina sphaerocephala and Verbesina fastigiata on germination and growth in Solanum lycopersicum and Cucumis sativus seedlings. Horticulturae. 8(7): 652. https://doi.org/10.3390/horticulturae8070652
Velasco-Ramírez, A.P., Virgen-Calleros, G., Hernández-Herrera, R.M., Velasco-Ramírez, A., Corona-Cervantes, M.P., Munguía- Lizalde, D.M. and Hernández-Pérez, A.R. 2021. Verbesina sphaerocephala extracts promote the development of Cucumis sativus and reduce the damage caused by Meloidogyne incognita. Mexican Journal of Phytopathology. 39(2): 339–353. https://doi.org/10.18781/r.mex.fit.2103-2
Velasco-Ramírez, A.P., Velasco-Ramírez, A., Hernández-Herrera, R.M., Murguia-Lizalde, D.M. and Velasco-Ramírez, S.F. 2022b. Extracts of Verbesina sphaerocephala and Verbesina fastigiata promote potted development of Fragaria ananassa. Agroindustrial Science. 12(1): 29–37. https://doi. org/10.17268/agroind.sci.2022.01.04
Velazquez-Gamboa, M.C., Rodríguez-Hernández, L., Abud- Archila, M., Gutiérrez-Miceli, F.A., Ruíz-Valdiviezo, V.M., Valdez-Salas, B., Luján-Hidalgo, M.C. 2024. Efecto de las fitonanopartículas de óxido de zinc y magnesio en la producción de rebaudiósidos y compuestos fenólicos en plantas de Stevia rebaudiana Bertoni. Biotecnia. 26, e2313. https://doi.org/10.18633/biotecnia.v26.2313
Vibrans, H. [accessed December 19, 2024.] 2010. Malezas de México. Available at: http://www.conabio.gob.mx/ malezasdemexico/2inicio/home-malezas-mexico.htm,
Viveros-Legorreta, J.L., Sarma, S.S.S., Guerrero-Zúñiga, L.A. and Rodríguez-Dorantes, A. 2018. Bioensayo del efecto de fenoles producidos por Myriophylum aquaticum en cultivo sobre Lactuca sativa. Hidrobiológica. 28(1): 109–119.
Xu, L. and Geelen, D. 2018. Developing biostimulants from agrofood and industrial by-products. Frontiers in Plant Science. 9: 1567. https://doi.org/10.3389/fpls.2018.01567
Yakhin, O.I., Lubyanov, A.A., Yakhin, I.A. and Brown, P.H. 2017. Biostimulants in plant science: A global perspective. Frontiers in Plant Science. 7: 2049. https://doi.org/10.3389/ fpls.2016.02049
Youssef, S.M., El-Serafy, R.S., Ghanem, K.Z., Elhakem, A. and Abdel Aal, A.A. 2022. Foliar spray or soil drench: Microalgae
application impacts on soil microbiology, morpho- physiological and biochemical responses, oil and fatty acid profiles of chia plants under alkaline stress. Biology. 11(12): 1844. https://doi.org/10.3390/biology11121844
Zhuang, X., Gao, J., Ma, A., Fu, S. and Zhuang, G. 2013. Bioactive molecules in soil ecosystems: Masters of the underground. International Journal of Molecular Sciences. 14(5): 8841– 8868. https://doi.org/10.3390/IJMS14058841
Zulfiqar, F., Casadesús, A., Brockman, H. and Munné-Bosch, S. 2020. An overview of plant-based natural biostimulants for sustainable horticulture with a particular focus on moringa leaf extracts. Plant Science. 295: 110194. https://doi. org/10.1016/j.plantsci.2019.110194