3.1. Mode of Action of Biofertilizer

The different mechanisms of action of biofertilizers, including nutrient uptake facilitation, phytohormone regulation, and phytoprotection, must be understood to effectively utilize their potential for increasing the ecological services of forest biomes and promoting production in agriculture sectors [32]. By supplying vital nutrients such as nitrogen, phosphorous, and potassium, biofertilizers preserve the soil’s natural qualities (Aloo et al. [81]; [Figure 2]). PGPMs are categorized into three major divisions: Arbuscular mycorrhizal fungi (AMF), plant growth-promoting rhizobacteria (PGPR), and nitrogen-fixing rhizobia [82]. PGPRs help plants grow and develop by making it easier for plants to take in nutrients (N₂ fixation and P solubilization), causing the root surface to grow (hormone production), or reducing the damage caused by pathogens, which is a sustainable way to increase crop yield [83]. Beneficial bacteria, including RHIZOBIUM [84], AZOTOBACTER [85], and AZOSPIRILLUM [86], have the capacity to fix atmospheric nitrogen into a form that is easily available for plant absorption and support plant development [Figure 3]. Phosphorus-solubilizing microorganisms, including PSEUDOMONAS and BACILLUS, may solubilize and mineralize both organic and inorganic phosphorus [87], making it available for plant absorption. In addition, several biofertilizers, such as AZOTOBACTER and AZOSPIRILLUM, release chemicals that promote root growth and aid in enhancing nutrient uptake [88,89]. Then, some biofertilizers, including AZOTOBACTER, AZOSPIRILLUM, and BACILLUS [90], release hormones that encourage plant growth, including auxins [91], gibberellins, and cytokinins. In addition to supporting root growth, blooming, and fruiting, these hormones assist in controlling plant growth and development.

Figure 2: The beneficial mechanisms of microbial strains as a biofertilizer and their role in maintaining soil fertility and enhancing crop productivity. [Click here to view]

Figure 3: Role of biofertilizers on physiological and biochemical properties of soil. [Click here to view]

In addition, biofertilizers can shield plants from a variety of biotic and abiotic stressors [92,93]. By producing antibiotics and other antifungal substances, for instance, they help reduce the severity of soil-borne infections [94]. They can also improve soil fertility and reduce environmental pollution by binding metals, which decrease the toxicity of heavy metals [95]. In addition, biofertilizers can encourage the formation of phenolic compounds, which are naturally occurring poisons that help protect plants from harmful organisms, which in turn can promote plant defense mechanisms[96].

3.2. Status of Biofertilizer Application in Agriculture and Forestry

In many parts of the world, particularly in developing countries, market momentum has increased due to concerns about sustainable agricultural and forest management, notably integrated nutrient management. As a result, the use of biofertilizers AMF, phosphorus-solubilizing bacteria (PSB), AZOSPIRILLUM, AZOTOBACTER, RHIZOBIUM, ACETOBACTER, seaweeds, etc.) has received more attention [97,98]. Demand for biofertilizers has unexpectedly increased in China, Canada, Argentina, and Europe, particularly in Spain, Italy, and Germany, as well as in the United States, India, and Africa [99,100]. These nations are putting a lot of effort into promoting the use and development of biofertilizers as a result of their growing demand for organic products and the realization of the great benefits that these fertilizers provide [101-103]. The global organic agricultural area is increased from 69.4 million hectares in 2017–74.9 million hectares in 2020, according to FiBL data [104]. The consumption of organic products, including biofertilizers, has increased along with the adoption of more organic farming techniques, which has enhanced soil fertility. However, plants get benefits when biofertilizers are used as seed or soil inoculants because they grow and take part in the nutrient cycle. In agriculture and forestry, biofertilizers are becoming more popular since they offer an alternative to chemical fertilizers, are less expensive, improve soil health, and increase production by 10–25% without harming the soil or the environment [105]. The use of biofertilizers can also help to reduce soil erosion and improve water retention [106], making it an attractive option for agriculture and forestry. Recently, research has focused on biofertilizers to improve soil fertility by reducing the salinization of soil [107], phytoremediation of metal-contaminated soils [108], and increasing the tolerance capacity to extreme events [109] to maximize agricultural yields. Some of the research has progressed from the laboratory to the field trial stage. Research has focused on the continuous improvement of technology to reduce costs, maintain quality, and increase the popularity of biofertilizers among farmers. New technology such as nanotechnology, introduced as nano-fertilizers, nano-biofertilizers, nano-pesticides, and nutrients, has aided in the development of advanced, low-cost, and environmentally friendly fertilizers [82]. To restore functional and resilient agricultural microbiomes in the modern era, high-throughput sequencing has been proposed as an adequate strategy to select the “missing pieces” (i.e., those microbes involved directly and/or indirectly in soil P cycling whose populations were disturbed or unbalanced due to unsustainable intensive agricultural practices) [110,111].

3.3. Application of Biofertilizer in Forestry and its Impacts

In forest ecosystems, bio-fertilizers can be an integral part of integrated nutrient management where nitrogen fixers, potassium and phosphorus solubilizers, growth-promoting rhizobacteria (PGPRs), endo- and ectomycorrhizal fungi (ECMF), cyanobacteria, and other beneficial microorganisms are used to improve nutrient and water uptake, plant growth, and plant tolerance to abiotic and biotic factors [9]. ECMF play a crucial role in the nutrient cycle in terrestrial ecosystems, particularly in forest ecosystems where ECMF forms a symbiotic relationship by forming a mantle and Hartig network of intercellular hyphae in the roots of forest species [112]. This relationship provides significant benefits for the restoration of forests and ecosystem soil aggregation and stabilization [113]. In addition, ECMF and mycorrhizal helper bacteria promote the growth of economically significant trees and cut fertilizer costs in an ecofriendly manner [114].

More than 7000 different species of fungi generate ectomycorrhizae [115], many of which are associated with significant commercial trees such as poplar, birch, oak, pine, and spruce [116]. ECMFs are symbionts with most temperate and boreal forest trees, providing soil nutrients and water in exchange for plant carbon, and they are also used to restore sites contaminated with heavy metals, affected by soil erosion, and degraded due to clear-cut logging and wildfire [117]. ECMF nursery inoculation enhanced the performance of conifers planted in oil sands reclamation regions on degraded gypsum soil [118]. Even ECMF appears to be particularly efficient in recovering heavy metal-contaminated forest soils [119]. Recent years have seen the discovery of many ECMF isolates and associated hosts with increased metal tolerance [120]. In tropical forests of New Caledonia that have been degraded by mining activity, the ECMF species Pisolithus albus has been employed as an inoculant of ectomycorrhizal endemic hosts (Acacia spirorbis and Eucalyptus globulus) to enhance plant growth and act as a protective barrier to hazardous metals [121].

In addition, ECMFs were used to restore clear-cut logging sites, and it was found that soil transfers from mature adjacent forests helped form ectomycorrhizas by getting rid of harmful rhizosphere bacteria. This makes the clear-cut habitat a good place for organisms that were already living there [122]. Moreover, depending on the intensity and frequency of fire, some ECMF species have the ability to sustain forest fires; these fungi may be important for bringing in new ectomycorrhizal plant hosts and making the soil more stable through leached nitrogen (ammonium (NH⁴⁺) and nitrate (NO^3- form) captured and transferred to remaining hosts in the post-fire area [123,124]. ECMF also has potential for use in restoring sites invaded by non-native plant species [125]. The invasion of pine trees into the Southern Hemisphere is one of the most widespread invasions of non-native species on Earth. It has turned native forests, grasslands, and shrublands into forests dominated by conifers and forced different ECMF communities to live together in their natural habitats [126]. However, pine invasions into native forests are always linked to a small number of non-native and invasive ECMF species taking over the roots of the pine [127]. Even though Pinaceae species spread quickly, they are often planted for forestry [128]. A different way to stop or slow the spread of pine hosts is to use non-invasive ECMF as inoculants for new plantations.

Plant growth is also affected by the make-up and activity of the associated bacterial community (phytomicrobiome), especially in the rhizosphere [129]. Plant-growth-promoting rhizobacteria (PGPR) can stimulate many direct and indirect mechanisms, such as phosphate solubilization, nitrogen fixation (N₂-fixing), and phytohormone production [130]. Early seedling establishment is difficult in semi-arid forests, particularly in oak (Q. brantii) forests in Western Iran, but the use of biochar and PGPR is effective in controlling water stress in oak seedlings [131]. Brevibacillus reuszeri MPT17 strain of PGPR helps Carya illinoinensis roots grow and boosts nutrient levels in plants and the soil around their roots, especially by making more phosphorus and potassium available [132]. Rhizosphere-promoting bacteria, such as Bacillus paramycoides JYZ-SD5, and the ectomycorrhizal fungus, Schizophyllum commune, both help Metasequoia glyptostroboides grow even when it is under a lot of stress from salt [133]. Furthermore, nitrogen (N^2-) fixation by moss-associated CYANOBACTERIA can promote moss growth and is an important source of new nitrogen in Eastern Canadian boreal forest ecosystems [134]. CYANOBACTERIA conduct massive amounts of nitrogen to forest ecosystems through nitrogen fixation, colonize feather mosses such as Hylocomium splendens and Pleurozium schreberi in the subalpine forests of Mt. Fuji [135].

Studies have shown that the use of biofertilizers in silviculture can increase tree growth, yield, and quality [136]. The AZOTOBACTER was found to be more effective at increasing organic matter and nitrogen, whereas the PSB treatment is more effective at increasing phosphorus and potassium. The PSB mobilizes restricted Phosphorus into a form that is available to the plants, which leads to the growth of seedlings [80]. As some examples, growth promoting bacteria isolates such as Lysinibacillus sphaericus, Paenibacillusquercus, Paramyrothecium roridum, and Lysinibacillus fusiformis displayed significant potentiality to increased growth of Eucalyptus pellita. The application of this biofertilizer to developing or recently grown Eucalyptus pellita in the field warrants further study. To determine whether the isolated strains from the consortia are successful at promoting plant growth in the cultivation of Eucalyptus pellita under pot trial conditions [137]. Forest biomes are rich in microbes and many aspects of the dynamics of microorganism values in their assemblages and their interactions with hosts remain unknown and require further investigation.

4.1. Formulation Complexity

The use of biofertilizers is receiving more attention and several of the products are currently offered globally [140]. In biofertilizers with living microbial cells, the formulation is one of the key elements that greatly influence the quality of the biological agents [35]. The preparation of inoculants is a critical multi-step procedure that involves one or more strains of microorganisms, an appropriate carrier, and additives that offer a safe habitat to protect them in challenging circumstances during storage, survival of strains, transportation, and establishment after introduction into soils. An efficient preface of microorganisms at the target region is provided by the right formulation, which also increases their activity once they have been injected into the host [141,142]. Another meaningful aspect is cost-effectiveness, which should be considered when selecting formulated products [143]. For commercial biofertilizer production, different carriers and organic matter should be available and cheap [144].

However, one of the key challenges for developing an improved formulation under field conditions is that a microorganism shows promising results under laboratory conditions. Manufacturers chose two or more types of microorganisms (e.g., RHIZOBIA and AMF, RHIZOBIA and PSB, diverse strains of AMF, or PSB) in a single product which help to maximize the resulting benefits for the host plants shown in several studies [39,71]. There are four types of biofertilizers, depending on the carrier’s physical characteristics and the material employed: formulations using solid carriers, liquid carriers, formulations with polymer entrapped carriers, and pressurized dry carriers [145].

Beneficial microorganisms’ metabolic activity quickly declines after production in liquid formulations [15,146,147]. Contrarily, solid formulations are problematic for non-sporulating bacteria because desiccation damages cell membranes, leading to cell death and loss of viability during rehydration, which has a significant negative impact on the commercialization of the product [148]. Despite the benefits of using the immobilized-cell formulation, it has still a limitation in large-scale production and field application because of the relatively high production cost [49,71,149,150], for the reason that the cost of polymeric carrier is high than the other methods [151]. Furthermore, cell mortality is one of the key difficulties of the bio-encapsulation process during the drying of encapsulated cells [150]. One of the major limits for polymer entrapped formulation is the survival of inoculums as low oxygen transfer [152].

Short shelf life and contamination are the key drawbacks of fluidized-bed dry formulation because of the moisture content of carrier-based inoculants [153]. In addition, safeguarding the finished product against infection is essential because unauthorized microorganisms have the power to completely alter the strain’s characteristics. However, it is essential to generate the strain in a sterile setting, which could raise production costs [154]. However, it is needed to improve formulations which are more effective, stable over time, better in quality, more consistent, cost-effective, and meet farmers’ needs.

4.2. Efficacy Changes with Climatic Conditions

The harsh and unpredictable environmental condition affects the efficacy of bio-formulation technology. Inoculants are subjected to biotic stress factors (such as microflora and microfauna) as well as abiotic stress conditions (such as soil pH, temperature, and salinity) during soil application [148]. In semi-arid environments, introduced inoculums struggle to survive inclement weather, including droughts, high salinity, inadequate irrigation, and even soil erosion, which quickly depletes the imported bacteria. In addition, bacteria have poorer tolerance for physical stress, particularly different temperature variations that occur during storage in peat carriers [152]. Due to their sensitivity to diverse environmental circumstances such as heat or drought, several biocontrol strains of bacteria are Gram-negative, which complicates their bioformulations [155]. Therefore, changing temperatures affect how adaptable and useful the microbial strains are: To solubilize Phosphorus (P) in pine forests under subfreezing temperatures, a PSB that is adapted to cold climates should be physiologically active. As a result of their functionality at room temperature, however, the psychrotolerant PSB species are likewise excellent options for P biofertilizers in any situation [156]. Nevertheless, the frequent use of plant growth promoting bacteria (PGPB) faces challenges during applications due to its inconsistent behavior and viability, particularly because PGPB cannot efficiently cover many plant species, its field application can generate inconsistent productivity, and the surrounding environment and microbial community can affect PGPB activity [83,157]. Finally, because of their poor growth in the culture system compared to their native habitat, many isolated PGPB species may find massive production difficult [158].

However, including giving due consideration to the rapidly changing global climate and region-specific forecasted conditions over the coming decades, which may affect product suitability and efficacy.

4.3. Storage Conditions

Microbial survival, carrier material qualities, biological effectiveness, and product storage life are all affected by storage conditions such as temperature, humidity, and sunlight intensity [159]. The carrier should maintain the viability of the microorganisms during storage in the farmer’s warehouse and have a lengthy shelf life and stability [150]. For the biofertilizer to be effective in the field, it must contain a sufficient number of viable cells during storage [160]. The shelf life of biofertilizers is another significant problem that can occur while using them. Live microbial cells used in biofertilizers have a limited shelf life (about 4–6 months under 20–25°C), and their storage and shipment require additional care and attention, increasing the cost of the product [110]. Biofertilizer should be stored properly, particularly at the correct temperature, to prevent the number of viable bacterial cells in the biofertilizer from falling below 108 cfu/mL. The lengthy shelf life of Acinetobacter baumannii, exceeding the required minimum of 108 cfu/mL at 6 months of storage, makes it a desirable biofertilizer product [161].

Biofertilizer should be kept cool, ideally in the refrigerator, to maintain their quality and shelf life. The recommended storage temperature for optimal efficiency and extended shelf life has been identified as 4°C [99]. The best storage temperature for Azotobacter venelandi NDD-CK-1 for up to 90 days was found to be 5°C, implying that inoculum can be stored at a low temperature for a prolonged shelf life [99]. On the other hand, Burkholderia spp. held at 28°C for 2 months grew more viable than those stored at 4°C, indicating that 28 °C is the best temperature for the studied rhizobia [160]. Whenever a 4°C inoculum is to be utilized, it must be cultured at 26°C for at least 7 days to commence microbial multiplication and achieve the requisite viable cell quantity. Biofertilizer stability and quality are very much dependent on storage conditions [162]. However, the above circumstance accounts for the product’s infrequent availability in rural locations [163].

4.4. Practicalities of Biofertilizer Application

Several methods of applying biofertilizers to soil exist, including root dipping, seed inoculation, and soil treatment using dry or liquid biofertilizers [29]. To create slurry for seed inoculation, carrier biofertilizers are diluted in water. To ensure that all of the inoculants are evenly distributed, sterile seeds are added to the slurry. The mixture is then air-dried before being planted. The root-dipping biofertilizer application technique is utilized for transplanted crops. Before being transplanted, the plant’s root is briefly submerged in a mixture of biofertilizer and water. Biofertilizer is applied as a foliar spray or as a soil application at a specific time when the farmer is prepared to plant the seed [164].

The use of biofertilizers is a biological strategy for the long-term sustainability of agriculture. However, there are significant obstacles to their use in enhancing agricultural output. As a result of biotic and abiotic stress, biofertilizers frequently perform less effectively in the field than they do in the laboratory or greenhouse [165]. Crops are grown in a variety of climates, soil types, levels of rainfall, crop varieties, and soil biodiversity. Because of these differences, the effectiveness of biofertilizers varies. In addition, because the inoculum needs time to colonize the root and increase its concentration, biofertilizers take longer to act than synthetic fertilizers [163]. The adoption of biofertilizers by farmers may be impacted by these reactions. The main issues with the use of biofertilizer include a lack of understanding within farmer communities of the value of microbial biofertilizer in terms of protecting the environment, inadequate encouragement, and promotion of the use of biofertilizer products by agricultural extension workers to farmers, a lack of acceptable carriers for the formulation of biofertilizer, and a lack of storage facilities [157,166]. In addition, because most biofertilizers are selective in their operations, the lack of labeling that indicates the expiration date and the identity of the microorganisms responsible for the biofertilizer’s formation may cast doubt on the validity of the materials [47].

4.5. Market Value

At present, people are focused on organic farming due to rising health concerns. As a result, consumers in developing countries are interested in biofertilizers, and the market for these products is expanding [167], as well as in developed countries such as Spain, Italy, and Germany, where there has been a surprising increase in biofertilizer demand [168]. The value of the worldwide biofertilizer market was $1.88 billion in 2022, and it is anticipated to reach $2.14 billion in 2023. With a compound annual growth rate (CAGR) of 13.2%, the biofertilizer market is projected to reach $3.51 billion in 2027 [169]. Legumes and N₂-fixing inoculants now dominate the global biofertilizer market [170]. According to the literature, rhizobia-based inoculants account for about 78% of the global biofertilizer market, whereas phosphate solubilizes and other bio inoculants account for 15% and 7%, respectively [167,170].

As per reports, India is the world’s fourth-largest user of Potassium bio inoculants, whereas the United States, China, and Brazil top the list in terms of the total consumption of these microbial products [171]. When looking at the market by geography, specifically by continents, North America dominates the global market for biofertilizers, closely followed by Europe in second place and Asia-Pacific in third place [172]. The remainder of the world has been grouped, with South America rising to take the top spot. The biofertilizer industry is expanding internationally as a result of the desire to boost food production sustainably [70].

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