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Sun, W.; Shahrajabian, M.H. Classification of Biostimulants. Encyclopedia. Available online: https://encyclopedia.pub/entry/49141 (accessed on 18 November 2024).
Sun W, Shahrajabian MH. Classification of Biostimulants. Encyclopedia. Available at: https://encyclopedia.pub/entry/49141. Accessed November 18, 2024.
Sun, Wenli, Mohamad Hesam Shahrajabian. "Classification of Biostimulants" Encyclopedia, https://encyclopedia.pub/entry/49141 (accessed November 18, 2024).
Sun, W., & Shahrajabian, M.H. (2023, September 14). Classification of Biostimulants. In Encyclopedia. https://encyclopedia.pub/entry/49141
Sun, Wenli and Mohamad Hesam Shahrajabian. "Classification of Biostimulants." Encyclopedia. Web. 14 September, 2023.
Classification of Biostimulants
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This entry is adapted from the peer-reviewed paper 10.3390/plants12173101

Biostimulants provide beneficial properties to plants by increasing plant metabolism, which promotes crop yield and improves the quality of crops; protecting plants against environmental stresses such as water shortage, soil salinization, and exposure to sub-optimal growth temperatures; and promoting plant growth via higher nutrient uptake. Biostimulants are classified as microbial, such as arbuscular mycorrhizae fungi (AMF), plant-growth-promoting rhizobacteria (PGPR), non-pathogenic fungi, protozoa, and nematodes, or non-microbial, such as seaweed extract, phosphite, humic acid, other inorganic salts, chitin and chitosan derivatives, protein hydrolysates and free amino acids, and complex organic materials.

biostimulant arbuscular mycorrhizal fungi microbial biostimulants pathogens plants abiotic stress tolerance

1. Introduction

Biostimulant application is known as an eco-friendly and novel farming practice and is relevant to two otherwise contrasting concepts, namely crop sustainability and intensification [1]. Biostimulant products already form a significant part of the global farming industry, indicating increasing trends over the years and in the future [2]. There are various reports regarding their positive impacts on crops, especially under biotic and abiotic stress conditions, and significant research is continuously conducted to find and/or produce new biostimulatory products, as well as to show the mechanisms of action behind the observed impacts. However, the variance in the composition of biostimulant products, as well as the lack of ordinary application protocols for the different products, may create inconsistencies between the observed results and complicate attempts to reveal the actual mechanisms behind the biostimulatory impacts, which may include physiological procedures, hormonal regulation, and morphological alterations. Biostimulants’ beneficial activities include the improvement of nutrient uptake, the induction of root growth, and the production of phytohormones; osmotic adjustment through the synthesis of organic osmolytes has also been confirmed. Biostimulants can also be applied to decrease the application of mineral inorganic fertilizer and are considered environmentally friendly tools with no significant negative impacts on fruit quality or total yield. Humic acids, fulvic acid, protein seaweed extracts, hydrolysates, N-containing compounds, botanicals, seaweed extracts, chitosan and other related biopolymers, beneficial bacteria and fungi, and inorganic compounds are the major categories of plant biostimulants. Modern crop production has to cope with abiotic and biotic stressors such as soil and irrigation water salinity, extreme and untimely weather phenomena, water limitations, infections from pathogens, and pests, which severely influence crop performance and the quality of the final products [3][4][5]. The most important advantages of biostimulants include improved profits, stimulated plant reactions, decreased operating costs, reduced application of fertilizers, improved root protection from soil pathogens, and enhanced drought tolerance; moreover, they repel pests, accelerate root establishment, boost fertilization, enhance stress tolerance, ameliorate fertilization, alleviate leaching, detoxify heavy metals and chemicals, and improve stomata opening and plant transpiration [3][4][5][6]. Biostimulatory compounds may also have positive effects on soil biology and are recognized as a good technique for recovering semi-arid areas and degraded ecosystems [6][7][8]. However, the variable composition of raw materials applied for the production of biostimulant products makes the task of revealing the mechanisms of action more difficult, and long-term research and standardization processes are needed [9]. Different sources of chitin and chitosan in nature are crustaceans (lobster, shrimp, king crab), fungi (Mucor rouxii, Penicillium chrysogenum, Aspergillus niger, Lactarius vellereus), insects (ladybug, wax worm, silk worm, butterfly), and mollusks (shell oysters, squid pen). Crustacean shells are the most notable chitin source, and chitin recovery involves three steps consisting of demineralization, deproteination, and the elimination of pigments and lipids [10][11][12]. Microbial proteases such as Lactobacillus sp., Bacillus sp., Pseudomonas sp., Serrati marcescens, etc., are the most significant strains applied in chitin and chitosan production [10].
Biostimulants containing organic substances, humic acids, amino acids, algae extracts, and carbon and boron increased plant growth, yield, and shelf life of onion bulbs [13], and the application of diluted honey extract (DHE) improved photosynthetic parameters, antioxidant activity, biomass production, and yield [14]. The use of seaweed extracts, vermicompost, and a mixture of animal waste increased yield and bulb traits [15]. Foliar application of vermicompost leachate, smoke-water, Ecklonia maxima extracts, and indole-3-butyric acid on seedlings of mustard greens grown in soils from goldmines boosted phytoremediation activities through the accumulation of heavy metals [16]. Foliar application of Kelpak SL and Asahi SL increased the nutritional value and improved the storage life of carrots [17], while root and foliar application of protein hydrolysates in lettuce plants grown under salinity conditions mitigated oxidative stress and increased glucosinolate and osmolyte content [18][19]. In intensive cropping sectors such as horticulture and floriculture, biostimulants can also boost nutrient use efficiency, partly substitute chemical fertilizer inputs, and ameliorate the quality and yield of crops [20][21]. Biostimulants based on microorganisms are a subgroup of the heterogeneous family of biostimulants, related to a microorganism (or mix of microorganisms) that can stimulate biochemical and physiological processes that benefit the nutrient efficiency, nutrient uptake, abiotic stress tolerance, crop quality, and/or yield of plants [22], which can moderately mitigate the damaging effects of intensive agriculture [23][24][25]. The most common microorganisms included in this group of biostimulatory products are the non-pathogenic and non-toxigenic bacteria of Azotobacter spp., Rhizobium spp., and Azospirillum spp., as well as different mycorrhizal fungi [24]. Mycorrhizas are a symbiotic association between fungi and plant roots and are present in several forms according to the fungal taxonomy and the host plant. Two important parameters that influence the distribution of these forms are the climatic and soil conditions and the host plant distribution [26][27] (Hart and Reader, 2002; Yang et al., 2012), and mycorrhiza can significantly boost the efficiency of mineral absorption, falling into two major categories: endotrophic and ectotrophic [28]. The main types of arbuscular mycorrhizal fungi (AMF) are related to the sub-phylum Glomeromycotina of the phylum Mucoromycota [29], and four orders of AMF, namely Glomerales, Paraglomerales, Archaeosporales, and Diversisporales, have been recognized in this sub-phylum, which contains 25 genera [30][31]. The protective mechanisms are credited to arbuscular mycorrhizal fungi-assisted alleviation of oxidative stress, rapid water uptake and nutrient absorption, and changes in the transcript levels of genes involved in signaling pathways or stress response [32][33][34], and the effectiveness of AMF is usually influenced by environmental variables and soil conditions [35].

2. Biostimulant Categories

Biostimulants are classified into two distinct groups based on their origin; one category includes products that have biological origins in pathogens or plants, and the second group consists of products that do not have biological origins [36][37][38]. Another classification approach divides biostimulant products into microbial biostimulants, which are obtained from arbuscular mycorrhizal fungi and plant-growth-promoting bacteria, and non-microbial biostimulants, which include plant micro-algae extracts, humic substances, and biopolymers such as chitosan [39][40][41][42][43]. Different compounds with bioactive properties can be used as biostimulants to boost plant growth and development under normal and stress conditions [44][45][46][47][48][49][50][51][52]. Salicylic acid is economical and quick in action, environmentally sound, and it also links with other elicitors to boost the biosynthesis of secondary metabolites [53][54][55]. Humic acid can increase plant growth, retain water, enrich nutrients, and suppress disease [56][57]. Fulvic acids are used in sustainable horticulture and can change plant primary and secondary metabolism and increase nutrient uptake, root growth, and crop tolerance to environmental stresses [58][59]. Protein hydrolysate biostimulants, mostly produced by chemical and enzymatic hydrolysis of plant- and animal-derived proteins, are based on a mixture of soluble amino acids and peptides and can increase the yield and quality of products as well as improve the nutrient uptake and abiotic stress tolerance of plants [60][61][62][63]. They are largely prepared from brown seaweeds, such as Ecklonia maxima, Ascophyllum nodosum, and Macrocystis pyrifera, and they include promoting hormones or trace elements such as Zn, Fe, Mn, and Cu [64][65]. Humic-like substances such as fulvic and humic acids may also show biostimulatory activity, since several reports have suggested improved crop performance attributed mainly to auxin- and cytokinin-like impacts; they are obtained from organic matter decomposition and metabolic products of soil microbes, and they have roles in plant growth via the improvement of soil physical–chemical properties and the boosted availability of nutrients in the rhizosphere [66][67][68]. The actual mechanisms of action seem to be the result of synergy between the several bioactive components in raw materials, although the impacts may change depending on the crop, soil type, and soil microbes present in the rhizosphere [69][70][71][72]. The most important impacts of chitin and its derivatives’ applications are that they stimulate and protect seed germination, stimulate stress resistance, mitigate negative impacts of abiotic stress, induce plant growth and development, improve soil properties and prevent nutrient leaching, improve the shelf-life of crops, chelate heavy metals, increase crop yield and quality, and protect against pests and pathogens, e.g., bacteria, viruses, fungi, insects, and nematodes [73][74][75]. Amino acids are the best candidates to boost stress tolerance through osmo-protection, ROS scavenging, metal chelation, and nutrient availability [76], which can notably impact the synthesis and stimulation of some enzymes and gene expression [77][78][79]. They can also be applied as signal molecules, like for inducing stomatal closure, as sensors of the nutrient contents of cells, or as regulators for inducing their own catabolism. Amino acids can manage the procedure of protein synthesis, strengthening plant growth, photosynthesis, and yield formation. They can increase nutrient assimilation, use, and translocation, as well as increase the quality of constituents [79]. Amino acids are well-known biostimulants due to their positive effects on yield and plant growth, and can mitigate injuries from abiotic stresses [80][81]. Amino acids also have a significant role in ammonium fixation and C4 metabolism and in the biosynthesis of different components, including isoflavonoids, flavonoids, cutin, aurones, sporopollenin, stilbenes, proanthocyanidins, suberin, lignins, catechins, phenylpropenes, lignans, acylated polyamines, and other different alkaloid derivatives. The largest and most diverse group of secondary metabolites in plants is phenols, which have good antioxidant effects and are involved in the regulation of photosynthesis, physiological activities, oxidation reduction procedures, and plant breathing [82]. Phenolic acids and their derivatives are coumarins, stilbenes, quinones, lignans, flavonoids, curcuminoids, and tannins, which have meaningful roles in plant development, especially in pigment and lignin biosynthesis, and of course, they have a significant role in protecting plants from stress [83].
Protein hydrolysate biostimulants, mostly produced by enzymatic and chemical hydrolysis of plant-derived and animal proteins, are based on a mixture of peptides and soluble amino acids, and can increase the quality and yield of products as well as the uptake and abiotic stress tolerance of plants [84]. Glomus, the largest and most common genus in the phylum Glomeromycota, forms symbiotic relationships with plant roots [85][86], which can boost the drought tolerance of the host plant, mediated by proteins with chaperone-like activity [87]. Trichoderma fungi have important functions in nature as plant growth promoters and antagonists of phytopathogenic fungi [88], and as rhizosphere inhabitants, they contribute to interactions with microorganisms, soil, arthropods, and plants at multiple trophic levels [89], and can be used as biocontrol and biopesticide agents [90]. Members of the genus Trichoderma are also used in different industry branches, like in the production of biofuel, antibiotics, and enzymes [91]. The main Trichoderma–plant interactions include their impacts on plant morphology, plant physiology, nutrient absorption and solubilization, disease resistance, yield improvement, and abiotic stress tolerance [92]. Trichoderma reesei is a genus of filamentous fungi and a superior cellulose source for industrial uses, and it can produce proteins, including different enzymes, cellulases, hemicellulases, and hydrophobins [93][94]. The endophytic fungus Heteroconium chaetospira can also penetrate through the outer epidermal cells of its host, pass into the inner cortex, and grow all over the cortical cells, consisting of those of the root tip region, without causing apparent pathogenic symptoms [95], and it can provide even more nitrogen to the plant than mineralizing plant-available organic nitrogen [96]. Arthrobacter species, which are Gram-positive chemoorganotrophs and obligate aerobes, are commonly identified among soil bacteria [97], being dominant aerobic bacteria under the class of families Micrococcaceae and Actinobacteria [98], and nutritional versatility is the principal feature of arthrobacters [99]. Acinetobacter spp. are Gram-negative coccobacilli that are aerobic, non-motile, and oxidative negative, with no glucose fermentation ability; they can be found in different environments [100] and can fix nitrogen, solubilize minerals, produce siderophores, and act as plant endophytes or epiphytes, which can help hosts in detaching pollutants and tolerating environmental stresses [101]. Moreover, the plant-growth-promoting traits of Actinobacteria entail phosphate solubilization, IAA, and siderophores [102]. They can also promote higher phosphorus content and plant growth and increase radical scavenging, plant phenolic components, and antioxidant activity [103]. Other important bacteria are Enterobacter spp., Pseudomonas spp., Ochrobactrum spp., Bacilus spp., and Rhodococcus spp. [104][105][106][107][108][109][110][111][112][113]. Figure 1 shows different classifications of plant biostimulants.
Plants 12 03101 g001
Figure 1. Different classifications of plant biostimulants.

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Subjects: Agriculture, Dairy & Animal Science
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Wenli Sun
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Mohamad Hesam Shahrajabian
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Update Date: 18 Sep 2023
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