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Review

Mitigating Drought Stress Effects in Arid and Semi-Arid Agro-Ecosystems through Bioirrigation Strategies—A Review

by
Gandhamanagenahalli A. Rajanna
1,*,
Archna Suman
2 and
Paramesha Venkatesh
3
1
ICAR-Directorate of Groundnut Research, Regional Research Station, Anathapur 515701, India
2
Division of Microbiology, ICAR-Indian Agricultural Research Institute, New Delhi 110012, India
3
ICAR-Central Coastal Agricultural Research Institute, Ela Goa 403402, India
*
Author to whom correspondence should be addressed.
Sustainability 2023, 15(4), 3542; https://doi.org/10.3390/su15043542
Submission received: 23 January 2023 / Revised: 8 February 2023 / Accepted: 10 February 2023 / Published: 15 February 2023
(This article belongs to the Section Sustainable Agriculture)

Abstract

:
Drought stress is most alarming and destructive among the abiotic stresses that increased in intensity in recent years affecting global food and nutritional security. The main resource limiting global agricultural productivity is water. The previous two decades have seen a surprising amount of study reports on genetically modifying plants to increase their ability to withstand drought, but actual progress has lagged behind expectations. Applying bioirrigation techniques in drought-prone areas might be a workable alternative strategy. It does apply to the usage of living things or biological creatures, such as potential microorganisms that can move soil moisture from a zone with enough water to plants that are drought-stressed through the modification of agricultural microclimate using agronomic strategies. Potential microorganisms include Gram+ and Gram− bacterial consortiums, as well as plant-growth-promoting rhizobacteria (PGPR). In addition to PGPR, the utilization of soil macro-fauna in agriculture, such as earthworms, lugworms, termites, etc., can be utilized and explored in the near future towards bioirrigation. Earthworms and other macro-fauna are abundant in soil, digging deep burrows in the sediment and providing aeration to the plants. PGPR evolved with plant roots to enhance plant resilience under biotic and abiotic stresses. Benthic microorganisms, which include bacteria and microalgae, for instance, have a 70–80% higher water-holding capacity. Through research findings, these benthic microorganisms can be successfully identified and used in agriculture, and they may prove to be a cutting-edge method to increase plant-water-use efficiency. Similarly, plant roots of legume plants act as bioindicators under drought-stress conditions. These new developments make a significant contribution to addressing the problems of food security that come with changing climate. This review article offers information on bioirrigation techniques, their potential, estimating techniques, etc. Overall, this article goes into detail about how bioirrigation techniques aid crop plants in overcoming drought stress. Future research should focus on creating the most appropriate and effective microbes, dealing with the problem of delivery systems, and evaluating potential organisms in the field for microbial formulations to improve plant performance under drought stress and significantly lower yield losses in drought-affected areas.

1. Introduction

Bio-irrigation [1,2,3] is a promising technique using microorganisms, bio-indicators or other organisms to promote the soil moisture/water movement in the soil matrix. The bio-geochemistry cycle and microbial community structure of the sediments inside the soil matrix are, nevertheless, significantly impacted by the applied irrigation or rainfall of the benthic macro- and micro-fauna. Therefore, bioirrigation techniques frequently depend on organisms referred to as “Ecosystem engineers” [4,5,6,7,8]. However, these ecological engineers alter the physical state of the sediment environments by improving the soil’s lining and the circumstances of other creatures present in the sediment environment [5,9,10]. These bio-indicators can occasionally act as an early-warning system for environmental changes brought about by irrigation [11,12]. Important organisms that contribute to the decomposition of soil include several bio-indicators, such as earthworms and springtails [10,12] (Table 1). Microflora and macrofloral species are used to alter the ecosystems in dry, semi-arid, and humid regions that are more susceptible to drought and increased water stress. However, large populations of burrowing and bio-irrigating macrofauna, such as arenicolidpolychaetes [13,14], thalassinidean shrimps and crabs [15], and Pygospio elegans, Claparede [16], also greatly alter the soil ecosystem. These bioindicators significantly change the rhizospheric zone surrounding the root surface. In this area, the plant and the microbes interact closely. Earthworms and other macrofauna have been employed for bioirrigation because they break up soil aggregates, reduce soil bulk density, increase soil aeration, and increase water infiltration (Table 1). These actions expose more area for microbial attacks [17]. By enhancing the physico-chemical and biological properties of the soil [4,8,18,19], earthworms are also able to improve the soil–plant nutrient supply, which increases plant productivity [20]. However, research on bioindicators or ecosystem engineers, such as lugworms, termites, and earthworms, has been mostly restricted to grasslands or environments with poor vegetation [14,21,22]. Numerous studies have examined earthworm reactions to tillage, chemical additives, temperature changes, and changes in land usage [23,24,25].
Rainfed agriculture, which accounts for 80% of all agriculture worldwide, plays a significant role in ensuring the world’s food security [26]. A total of 90% of the rainfed agriculture is in Latin America, 95% in Sub-Saharan Africa, 75% in East and North Africa, 65% in East Asia, and 60% in South Asia [27]. Climate change, water scarcity, and population growth all pose threats to rainfed farming by making it more susceptible to droughts and other extreme weather conditions. For thousands of years, India’s expanding population has relied heavily on agriculture. In India, rainfed agriculture sustains around 61% of the nation’s farmers and comprises 55% of the net sown land area [28]. With an area of 86 Mha and a production rate of 40% of the world’s food grains, India is the world leader in rainfed agriculture [26,29]. Still, the intensification of agriculture in India and other agriculturally dominant nations of the world led to climate change and frequently caused higher and more frequent droughts and floods. In addition to these, the rainfed production system is seriously threatened by soil degradation, low soil fertility, low soil carbon content, and widespread micronutrient shortages. Therefore, the world demand of water supplies is rising, and agriculture is becoming more and more dependent on water. The most precious input, water, has a significant impact on agricultural productivity, especially in semi-arid regions [30]. Rainfall that is above average results in runoff and leaves crop plants without access to moisture unless in situ moisture conservation is undertaken. The primary abiotic stress that lowers agricultural yield globally is drought, which affects plant growth on around 45% of the net cultivated area [31,32]. With the rising intensity of land use, improper water management is becoming an increasingly serious issue [33,34]. Drought stress has a deleterious effect on plant growth, especially leaf area and dry matter accumulation have been reduced, which may ultimately cause plant mortality. By regulating the soil fauna’s abiotic habitat by irrigation, the risk of anhydrobiosis and dehydration is reduced. Drought stress may have a significant impact on grain quality and yield through the reduced growth and development of plants. Plants show signs of drought stress, such as leaf rolling, yellowing of leaves, and irreversible wilting, as the roots grow to absorb more water. In areas that are prone to drought, irrigation is utilized to make up for the soil-water deficit and maintain pasture growth [33]. Therefore, new things like bioirrigation help a lot with the problem of food security in a world where the climate is changing. This review article talks about their potential, how to estimate them, etc. Overall, this article gives a lot of information about how bioirrigation helps crop plants deal with the stress of drought.
Studies on grassland, forests, and the ocean have revealed that macro- and micro-fauna under non-agricultural vegetation can be used to detect moisture stress [20,35,36]. However, in terms of agricultural crops, bioirrigation has not yet been used in India or the rest of the world to retrieve plant biophysical, chemical, physiological, and economic characteristics. However, microorganisms like rhizobactiraia have been used extensively under field conditions to impart drought tolerance. Yildirim et al. [37] explored a biological strategy employing plant growth and promoting rhizobacteria (PGPR) inoculation in crop cultivation. The negative consequences of one or more environmental stresses can be avoided by using PGPR and AMF in the fingermillet + pigeonpea system [3], maize + pigeonpea system [38], maize + faba bean system [39], and in the maize-wheat system [40]. Additionally, the discovery, selection, and use of appropriate beneficial microbes can expand the choices of resolving environmental issues [41,42]. Utilizing bio-indicators can help identify the root of an environmental issue and serve as an early warning system for environmental changes [43]. However, there is not a lot of material on agriculture crops from regarding drought stress by bio-irrigation. Focused future study is required to identify the appropriate bacteria and address the issue of delivery mechanisms and field evaluation of potential organisms [44].
Table 1. Indicators used in bioirrigation.
Table 1. Indicators used in bioirrigation.
Bioirrigation Agent/OrganismCrop/ConditionOutcomeReference
Lugworms (Arenicola marina)Germany under sandy tidal bayLugworm activities prevent the sediments from developing sand to mud successively[5]
Lugworms (Arenicola marina)Strong bioindicator in coastal sediments-[36]
Diptera insect, Chironomusplumosus (Chironomidae)Soil experiments2.5 times higher sediment respiration;
widespread and abundant bioirrigator, building deep burrows in the sediment and pumping water through the burrow
[2]
Earthworms, Aporrectodea caliginosa; Lumbricus rubellus and Aporrectodea longaEffects of irrigation and effluent distribution on the diversity of earthworm species in pasturesSheep farms have larger earthworm populations and biomasses than dairy farms[7]
Mites, Acariformes (Arachnida; Acari)
Springtails (Arthropoda: Collembola)
Irrigated and non-irrigated conditions with treated municipal wastewater and dairy parlour washings on SRC willow plantationsThe amount of earthworms and mites in the SRC willow plantation was considerably impacted by previous land use, with an increased abundance of earthworms under previously planted grasslands[12]
Rao et al. [36] used lugworm (Arenicola marina), which has deep burrowing effect on oxygen, calcium, and nutrient content along with pH and alkalinity in porewater profiles and sulphide in sediments, to study the role of macrofauna in coastal sediments. Reduced levels of biological yield, essential oil percentage, essence yield, and chlorophyll a and b are all effects on drought stress. By using humic acid PGPR drivers in stress-level drought, the plant becomes more stable and exhibits the desired performance characteristics [45]. Similarly, Chironomus plumosus (Chironomidae), a Diptera insect, was employed as a bioirrigator in soil tests. It was shown that Chironomus plumosus is a ubiquitous and abundant bioirrigator, digging deep burrows in the sediment and pumping water through the burrow [2].

1.1. Need of Bioirrigation in Agriculture

Drought stress is considered the most harmful abiotic stress, it affects global food security, and has become worse over the past few decades. Drought stress affects plant–water interactions at the cellular and systemic levels, which results in economic losses in agriculture [46]. Numerous studies have examined how drought stress affects plant growth and soil indicators [29,44,47]. Abiotic stress, which includes drought, is one of the most significant problems affecting plant growth and development and altering agricultural needs, but plants are continually vulnerable to it. Around the world, intensive research is being conducted to develop drought-tolerant genotypes/varieties, alter crop calendars, establish resource-management strategies, etc., to deal with drought stress [26,48,49]. The majority of these technologies are expensive and highly variable under field conditions. According to recent studies, microbes can assist plants in coping with drought stress [44]. Rhizobacteria that promote plant development (PGPR) can survive in harsh conditions and protect plants from the damaging effects of environmental stresses [50,51,52]. Although a sizable number of research papers on improving drought tolerance in plants by genetic modification have been published, the pace of the development has lagged behind expectations. By utilizing rhizospheric bacteria that have evolved along with plant roots in harsh environments and that have adaptive features that improve plant vigor in the face of biotic and abiotic stresses, we propose a workable alternative approach.
Crops’ drought tolerance has been increased using a range of techniques, including classic selection techniques and genetic engineering [53]. Higher plants respond to a water shortage by changing their morphology and biology through stomatal closure and osmotic adjustment. Together, these adjustments increase water efficiency, water conservation, and soil moisture extraction [54,55,56,57]. Reactive oxygen species (ROS) production through free radical’s formation linked to membrane damage and lipid peroxides accumulation, can happen in highly water-stressed plants despite adaptations [58,59,60]. ROS production in plants is directly linked to stress, which causes membrane damage and stress signaling [61]. In chloroplast thylakoids, ROS are made in the reaction centers of photosystem I (PSI) and photosystem II (PSII) [62]. Several enzymes take part in redox reactions that can any of the ROS can process [63].

1.2. Bioirrigation Potential in Agricultural Crops

In India and other arid- and semi-arid regions of the world, drought stress is distinct from other abiotic stresses in that it lowers the output of cereal, pulse, and oilseed crops. Numerous attempts have been made to decrease the effects of drought stress on growth and productivity because the restriction of plant growth and productivity caused by drought stress is particularly harmful. As a crop with deep roots, pigeonpea is referred to as “Biological plough”, and it nodulates freely. Pigeonpea is commonly grown in India and other developing countries in Asia, Africa, and Latin America because it can often withstand drought. It is known to have “induced systemic tolerance” (IST) when microorganisms alter the physical and chemical makeup of plants to increase their tolerance to drought [44].
Using microorganisms for bioirrigation can boost production and productivity by improving the hydraulic lift of soil moisture from deeper soil profiles to impart drought stress tolerance. According to Caldwell et al. [64], the water that plant roots absorb from the soil is intermittently released back into the atmosphere. The same processes that controls water input into roots also controls water outflow from roots. A potential water gradient is produced by the transpiration of water loss from the leaves and flows from the soil to the root xylem, which is where water influx occurs. In these circumstances, water might exit the roots and enter the soil along a potential water gradient that has been reversed. This will make it easier to determine which hydraulic lift bioirrigation techniques work best when there is drought stress.
Concurrently, plant-growth-promoting rhizobacteria (PGPR) use has multiple advantages in the form of nitrogen fixation, releasing plant-growth-promoting hormones (phytohormones) and imparting drought-stress tolerance by releasing polysaccharides. According to Zahedi and Abbasi [42], PGPR application had a more positive impact on phytohormones that promote growth compared to control treatment in Iran. Concurrently, the impact of conventional irrigation using PGPR has a direct correlation with enhanced soil humic acid content [65]. Abscisic acid accumulation was decreased by using PGPR. When PGPR was applied to stressed soybean plants, the buildup of polyamines was shown to be significantly reduced. Therefore, PGPR can be effectively utilized with hydraulic lift mechanism plants to minimize the harm caused by drought stress. However, the impact of microbial activity on agricultural crops has not received much attention in Indian agriculture. Moreover, no similar research in agricultural crops has been recorded, whereas the possibility of bioirrigation has been investigated using different bio-indicators.
Earthworms are capable of enhancing the soil’s physical, chemical, and biological parameters as ecosystem engineers [18]. By improving the soil’s capacity to deliver a variety of ecosystem services, such as plant nutrition availability and earthworms that can boost plant productivity [20,66] However, most of the bioirrigation studies using earthworms confined to pasture, forage crops, and forest tress. However, these studies should be carried out in agricultural crops as earthworm and other insect populations vary in cultivated conditions. Since earthworm populations would be much higher under natural systems, as compared to cultivated conditions where death of earthworms occur at a faster rates [67]. Therefore, the benefits of the earthworm’s introduction to cultivated arable soils are higher under no-till/zero tillage/conservation agriculture conditions and in permanent pasture crops [68]. Providing moisture and effluent application significantly influences earthworm communities through improving the reproduction of adult earthworms, and also the cocoon survival rates [69]. By changing the equilibrium of the reaction rates in soil matrix and solute transport, macrofaunal activity has a significant impact on carbon and nitrogen cycling, as well as on the solute fluxes across the soil-matrix–water interface [70]. The removal of reduced metabolites and the provision of organic substrates and terminal electron acceptors for microbial metabolism via bioirrigation have a significant impact on the permeability, stability, and composition of sediments [36,70,71]. Therefore, earthworms could be promising bio-indicators in drought-stress regions.

1.3. Cropping Systems as Bioirrigation

Hydraulic lift and redistribution, a mechanism used by many herbs, grasses, shrubs, and trees, involves lifting water via roots from the subsoil and redistributing it to the topsoil [72,73]. Hydraulic lift refers to the movement of water via a plant’s root system across soil layers with different water potentials [64,66,74]. Water is passively lifted by the roots due to stomatal closure and leaf transpiration. If water is transferred into the topsoil, plants with shallow-root systems associated with deep-root plants may have access to enough moisture [64,66]. Through the differential root systems of plants, the hydraulic lift is the mechanism used to transfer water between soil layers by adjusting the soil-water potential. By including deep-rooted plants in intercropping systems, it may be possible for the associated crops to use hydraulic lift to obtain water from the uppermost soil layers [66,75]. However, identifying hydraulic lift plants is difficult under filed conditions and needs isotope studies. Sekia and Yano [75] studied the hydraulic lift in the pigeonpea–maize association in a glasshouse experiment using deuterium/hydrogen isotope (D2O). They also explained the isotope mechanism injection into the plants. They discovered that the maize plants’ xylem fluids had a greater deuterium concentration than expected, confirming that pigeonpea took water from deeper soil layers and moved it into the topsoil, thus enhancing microbial activities.
Saharan et al. [76] and Singh et al. [3] conducted field studies on bioirrigation in the presence of a PGPR, arbuscular mycorrhizal fungi (AMF), and common mycorrhizal network (CMN) effects between two intercropping species in finger millet and pigeonpea systems. Saharan et al. [76] found that when drought circumstances exist and there is “biofertilization” with PGPR and AMF in the pigeonpea + fingermillet system, pigeonpea, a deep-rooted legume, can benefit from finger millet (shallow-root cereal). The results showed that the CMN connecting pigeonpea and finger millet exerts a clear positive influence in this simulated intercropping system by reducing the negative effects of drought conditions on finger millet. Similarly, Sharma and Guled [77] obtained the increased water-use efficiency of 2.76 kg ha-mm−1 in a pigeonpea + sesame (1:2) intercropping system in set-furrow plots with vermicompost applied at 2.5 t ha−1; however, they were unable to identify the cause of the higher water-use efficiency. However, their findings did not take into account intermediate moisture levels, and to our knowledge, there is no literature available on bioirrigation in India.

1.4. Beneficial Microorganisms as Bioirrigation

As of today, microorganisms are being utilized or exploited as a source of nutrients to increase the productivity of cereal, pulse, and oilseed crops in systems of intercropping and succession planting. These microbes fix atmospheric nitrogen through their symbiotic nature, thus reducing the application of excess chemical fertilizers in pulse and oil seed crops. Besides N fixations, some studies reported plants having higher drought-tolerance capacity in dryland crops when PGPR are applied. Research studies show that plant inoculation with bacteria from abrasive habitats increased drought-stressed plants’ survival rates, as well as their capacity for photosynthesis and produce biomass [78]. However, some studies found that PGPR imparts drought tolerance in plants by releasing polysaccharide (Table 2). The studies of Timmusk [79]; Kim et al. [80] highlighted that free-living PGPR were found to be a promising broad-spectrum alternative technique to enhancing plant development under drought-stressed plants. Beneficial microbes, such as Bacillus spp. [46,78], PGPR [81], AM fungi [82], and alginate, a hygroscopic bacterial polysaccharide [78], are used in maize and wheat for increased plant tolerance. These microbes, especially the one belonging to the Bacillus genus, survive under the low-water potential of −0.73 MPa conditions, and provide tolerance to plants due to reduced ascorbate peroxidase, catalase, and glutathione peroxidase activity [46]. Concurrently, the inoculation of maize plants with Bacillus spp. enhanced leaf-water potential, leaf-relative water content, and decreased leaf-water loss.
Gram-positive rhizosphere bacteria are commonly used due to their enhanced ability to colonize under stressful environmental conditions via creating endospores [83,84]. By creating phytohormones such as IAA and cytokinins the addition of PGPR promotes root development through a higher number of root tips and an enhanced root surface area [46,85]. Proteins like dehydrins are produced by plants to combat abiotic stresses like water scarcity [86]. Soil microbes, such as AMF and PGPR, can build a common mycorrhizal network (CMN) that extends beyond plant roots and facilitates long-distance nutrient mobilization, and water transfer may also be helpful [39,40,87]. It is possible to accomplish sustainable intensification through intercropping with a variety of crops, but doing so necessitates the fusion of several fields, including agronomy, soil, microbiological, and social sciences [88].
Sustainable agricultural production is possible in most cases when chemical fertilizers are used in conjunction with biofertilizers [89]. Among these PGPR, species belonging to the Azospirillum and Azotobacter genus have garnered attention as potential biological fertilizer producers due to their widespread geographical distribution, host plants, and the wide range of significant crop plants that they can interact with synergistically, including rice, wheat, maize, sorghum, and sugarcane [90]. Whereas, these bioinoculants could also induce systemic changes in regulating the phytohormones production, as well as by making alterations to the root hydraulic properties by the induction of aquaporins in the plasma membrane [91,92,93]. Biofertilizers have widely been used as a source of nutrients; however, there is a lack of research regarding their use for improving water-use efficiency. To determine the plant physiological status under drought, future research aims are to estimate the hydraulic lift and its associated contribution to the neighboring intercrops (Figure 1). This could have significant effects on how crop stress is detected, how agricultural fields are managed, and notably, how to find the best microbes that use nutrients and water efficiently.
Table 2. Different mechanisms lead to different effects of PGPR variants on crops.
Table 2. Different mechanisms lead to different effects of PGPR variants on crops.
RegionCrop and ConditionPGPR TypeMechanismObservationReference
IndiaMustard
Field study
Bacillus casamancensis, Bacillus sp. MRD-17, and Bacillus aryabhattai NSRSSS-1Extracellular polymeric substances (EPS)B. aryabhattai strain NSRSSS-1 demonstrated the maximum chlorophyll-a content after inoculation[94]
IndiaMustard
Pot experiment
Bacillus sp. strain MR D17 & Bacillus cereus strain NA D7Significantly increased DREB2 and DREB1-2 gene expression; essential function for ABA-independent pathway in improving toleranceProline, amino acids, and sugar osmolyte accumulation is increased, which improves osmotic adjustment and RWC in the leaf tissues[47]
IndiaMaize
Pot/Lab experiment
Bacillus amyloliquefaciens, Bacillus licheniformisProline, carbohydrates, and free amino acids were all enhanced by osmoregulation, while electrolyte leakage was minimisedSugar levels increased, and seedlings exhibited physiological reactions to counteract the deleterious consequences of drought stress[46]
Sweden and EstoniaWheat
Pot/Lab experiment
Bacillus thuringiensis;
Paenibacillus polymyxa
Biofilm of rhizosphere bacteria
has protective role against drought stress. Volatiles emitted from the inoculated plants can provide stress resistance.
Changed root hairiness; higher net assimilation rate[78]
ArgentinaMaizePot/lab experimentAzospirillum lipoferumGibberellins elevated ABA levels and reduced the effects of drought stressMaize growth enhanced[95]
Republic of KoreaSoybean
Pot/lab experiment
Pseudomonas putida H-2–3Reprogramming the expression of chlorophyll, stress hormones, and antioxidantsPlant growth was enhanced by P. putida’s secretion of gibberellins.[96]
PakistanWheat
Pot/lab experiment
Rhizobium leguminosarum (LR-30), Mesorhizobium ciceri (CR-30 and CR-39), and Rhizobium phaseoli (MR-2)Wheat growth and drought tolerance index were all improved by IAA produced by the consortiumsIncreased the seedlings’ root length to lessen the impact of the drought[97]
ArgentinaMaize
Pot/lab experiment
Azospirillum sp. and
Herbaspirillum sp.
Lower ZmVP14 gene expression, which is essential for the production of abscisic acidIncreased biomass production, higher amounts of carbon, nitrogen, and chlorophyll, and decreased levels of ethylene and abscisic acid[98]
CanadaPisum sativum
Pot Exp.
Variovorax paradoxus 5C-2The enhanced increase in xylem abscisic acidSymbiotic nitrogen-fixing bacteria boosted nodulation, preventing a drought-induced decline in nodulation and seed nitrogen content[93]
ChinaPisum sativum
Pot Exp.
Variovorax paradoxus 5C-2ACC) deaminase (ACCd)-containing rhizobacteriumEnhanced shoot-to-root xylem fluxes of K by 2.1 times and by 1.8 times, respectively.[99]
ChinaAlfalfaSinorhizobium meliloti 1021-Engineered strainCytokinin (ipt gene) excess productionIncreased zeatin, nitrogenase, and antioxidant enzymes in leaves[100]
USASquashGlomus intraradicesActive aquaporins and AM colonisationUnder drought, the host plant’s root hydraulic conductivity (Lpr) increases or decreases[91]
SpainPhaseolus vulgarisGlomus intraradicesHigher root plasma membrane aquaporins and root hydraulic characteristics (PIP)Protein abundance, PIP gene expression, and root hydraulic conductance (L)[92]

1.5. Methods to Analyze the Effect of Bioirrigation Potential

Existing quantitative models of bioirrigation mostly rely on chemical, rather than ecological, information, and the depth dependence of bioirrigation activity is either imposed or modified using a data-fitting technique. With the help of stochastic simulations of 3D burrow networks, some studies by Lamande et al. [101], Blouin et al. [102], and Hallam et al. [67,103] estimated the potential of bioindicators by examining the volumes and wall surface areas of burrows, as well as their variabilities, which are calculated as functions of sediment depth. They observed that bioirrigation coefficients for O2 are greater than those based on vertical pore water chemical profiles. They also estimated based on the carbon dioxide emission that is produced when soil organisms respire, and also assessed based on soil respiration. In general, plant roots, the rhizosphere, microorganisms, and animals all respire in the soil. Koretsky et al. [1] observed a 2.5-fold increase in sediment respiration through Chironomid sprescens organism when compared to sediment without organisms. Likewise, Baranov et al. [2] identified a new and promising technique for measuring sediment ambient respiration is the ‘Raz respiration assay’. They observed that, in direct proportion to the level of bioirrigation, sediment respiration is rising.
Timmusk et al. [78] employed a GC-MS analysis to measure the benefit of bacterially inoculated plants by measuring the emission of terpenoid and benzenoid compounds from wheat leaves, including a-pinene, limonene, a-phellandrene, and camphene. They also kept track of the emissions of volatile organic compounds (VOCs), which they discovered to be a viable tool for determining how well-adapted various bacterial strains are to drought stress.
The bioirrigation potential or mechanism under field condition through crops or by the organism is not known. Under the intercropping system, quantifying the yield from base and inter crops is the existing method of understanding the effect of bioirrigation and biofertilization. However, the sensor-based instrument called ‘Dynagage’ has been developed by Dynamax Inc, VELCRO, which is a trademark of Velcro USA and is a Sap Flow Sensor, can be used to measure the moisture movement with the plant. Therefore, there is need to develop techniques and methods to identify bioirrigation potential under field conditions.

1.6. Bioindicators Effect on Soil’s Physical and Chemical Properties

Bioindicators like earthworms, lugworms, termites, and active plant roots greatly influenced the soil’s physical and chemical properties. Among these bioindicators, earthworms occupy the prime position in modifying soil structure through their burrowing nature. Research studies reported that earthworms improved soil quality through higher soil organic carbon and organic matter [67,102,103] under pasture fields. Thus, earthworms directly affect soil structure by generating pores of various sizes, branching, and sinuosity, all of which have an effect on the soil’s ability to store water [104,105]. These earthworms improve soil structure by aggregating, resulting in soil macropores of >1 mm, which significantly increased water flow [67,106]. As a result, the soil’s micropores and aggregation were improved, increasing soil-water retention. Concurrently, Hallam et al. [67] observed a higher water-holding capacity (~9%), water that is readily available to plants (~21%), and a lessened soil compaction (6% decrease in bulk density). Previous studies from Lamande et al. [101]; Blouin et al. [102]; Hallam et al. [67] claimed that earthworms had a beneficial effect on water flow. Earthworms (Allolobophora chlorotica) can regulate soil-water flow and have a substantial impact on soil hydraulic parameters by vertically burrowing [103]. Earthworms significantly increase plant-water availability, field-water content, and water-storage capacity [107]. Earthworms in the soil will significantly lessen infiltration during excess floods, reducing the adverse implications of flooding [67]. As a result, these improved the soil’s physical conditions and had a favorable effect on plant development. According to Hallam et al. [67], van Groenigen et al. [108], and Hallam et al. [103], earthworms greatly increase the growth of wheat plants and the dry biomass of grass-clover shoots by 58%. Along with earthworms, benthic macrofauna such as lugworms and Arenicola have a favorable impact on changing the soil’s biogeochemistry, which in turn affects plant development. Although termites’ impact on hydrology has been recommended as a restoration strategy, it is neither universal nor fully understood.
The interaction between lugworms and the sediments revealed that the fine sand area’s lugworm exclusion plots had the maximum capacity of tying together small particles in the top sediment layer [5,71]. Lugworm’s presence drastically decreased the surface sediment’s chlorophyll concentration. The presence of Arenicola had the greatest effects on chlorophyll concentration during the months with the maximum feeding activity. As a result, the amount of chlorophyll in the sediment appeared to be significantly influenced by lugworm-feeding activity. By establishing pores of various sizes and stabilizing macro-aggregates [109], plant roots and the related mycorrhizal fungus help to enhance the structure of the soil. The increased micropore volume in the roots of crops with thick and fine roots, such as grass [110], clover [111], and pigeonpea [3], increased the amount of water available to plants [112,113]. However, there is a need to establish a relationship between earthworms, lugworms and termites with associated root system under field conditions.

1.7. Bioindicators’ Effect on Biological Properties

The organization of the microbial population is influenced by bioirrigation [114,115]. Because biofilms include sugars and oligo- and polysaccharides, they are particularly helpful for plant growth and development. As a result, there is more water available in the root medium [78]. In fact, studies have demonstrated that polysaccharide alginate, even in small amounts, aids in preserving a moist micro-environment in the biofilm (Chang et al., 2007). The bacterial polysaccharide alginate, which is hygroscopic, can be crucial in determining a biofilm’s ability to improve a seedling’s hydration status [116]. Bacterial alginate can hold a lot of water and release it gradually [117,118], keeping root cells moist for long enough to allow the cellular metabolic adjustments needed to increase drought-stress tolerance. Alginate’s increased ability to withstand droughts may be attributed to its hygroscopic qualities, but it may also be a result of its participation in biofilm construction, which lowers evaporation loss. The anatomical features of the leaflets were modified by the combined use of B. subtilis and B. amyloliquefaciens, with a rise in the thickness of the upper epidermis, lower epidermis, palisade tissue, spongy tissue, and vascular bundles in tomatos [119]. More than 80% of rhizosphere microbiomes, including Bacillus sp. and Enterobacter sp. [120,121], excrete the phytohormone indole-3-acetic acid (IAA) that controls various plant physiological processes, such as growth, division, differentiation, and gene regulation, as well as responses to environmental variations such as light and gravity [121].

1.8. Feasibility of Bioirrigation in Dryland Agriculture

Drylands are known for their lack of precipitation, low soil organic matter, susceptibility to erosion, desertification, deficit soil moisture, loss of fertility and salinization [29,122]. Therefore, technologies that improve water-holding capacity along with the fertilizer-supply mechanism a best suited for increasing the production potential of dryland regions compared to individual approach-based strategies. According to Gopinath et al. [29], controlling soil and crop water is crucial for enhancing productivity and bridging yield gaps in rainfed regions. In such cases, bioirrigation techniques combined with modified agronomic practices are crucial in increasing crop productivity. A potential bioirrigation method should be for bioindicators to maintain moisture on the body surface for a long time and then transport it to nearby plants during drought stress (Figure 1). This will significantly lessen the effects of the drought stress by increasing moisture intake, and subsequently, improving yields for dryland crops. However, the choice of cropping systems should include deep-rooted crops (pulses) as intercrops and shallow-rooted crops as base crops (millets and coarse cereals). According to studies by Cohen et al. [95], Vardharajula et al. [46], Timmusk et al. [78], Bandeppa et al. [47], and Vikram et al. [94], the current microbial formulations, such as PGPR-including species belonging to Bacillus, are known to improve plant drought tolerance by releasing stress tolerance compounds. Together, bioirrigation and the drought tolerance mechanism have a greater advantage in dryland areas. Deep-rooted plants, such as pigeonpea, participate in the hydraulic shift process during drought stress, where water movement could be coming from the deeper soil layers. Then the seed-treated microorganisms should break down in the soil, hold moisture, and transport it to the neighboring plants. However, these assumptions need a research base for the feasibility of bioirrigation in dryland regions.

1.9. Opportunities for Bioirrigation Potential

  • There is a need to isolate benthic zone microbes, which are having higher water-holding capacity over the existing microbes, which are being utilized only for nutrient purpose;
  • There is a need to identify and develop suitable microorganisms that have a higher bioirrigation potential to mitigate the drought stress in major cropping systems;
  • Some available microbial inoculates are tolerant to harsh environments and there is a need to identify the method of application with the rapid enhancement of plant stress tolerance with higher water-holding capacity;
  • There is a need for extensive studies on using beneficial microorganisms with the judicious use of available irrigation water under variable tillage conditions, which can lead to the production of eco-friendly cultivation practices;
  • This enlists new possible options for using earthworms, termites, lugworms, etc., under diversified cropping systems in conjunction with microbes;
  • They are wider options for breeders to breed efficient drought stress cultivars with efficient rooting patterns to extract moisture from deeper layers.

2. Conclusions

Rainfed regions provide the majority of the food for poor populations in under-developed and developing countries, despite geographical variances in rainfed agriculture. Rainfed areas are crucial for ensuring global food security. Due to the unpredictable nature of rainfall, increasing frequency of mid-season droughts, intense rainfall, and their increasing frequency over the past few years, rainfed farmers are becoming more and more vulnerable. Crop yields are low in these conditions, leaving large production gaps in rainfed regions. Bioirrigation may be a technology that significantly enhances crop yields. An integrated strategy that effectively utilizes bioirrigation resources and suitable soil management practices is required to boost the profitability and sustainability of rainfed farming in these areas. Microbes like bacteria, fungi and actinomycetes are bioindicators that have been found to be promising in developing drought tolerance in crops under rainfed regions. However, there is no information available, though, about how these bacteria contribute to the bioirrigation potential. Earthworms, lugworms, termites, and other bioindicators found in the soil have been shown to demonstrate aeration at the root zone of crops and could be used as bioindicators. The findings suggest that aerating the root zone can improve yield and quality by altering the composition of the soil’s microbial community, leading to greater nitrogen-fixing and potassium-solubilizing microorganisms in plants’ root zones. More research is required in this special field of utilizing bioirrigation to mitigate the effects of drought stress. To increase productivity in rainfed regions all over the world, it is crucial to identify the appropriate bioindicators for specific regions. In order to maximize the potential of rainfed agriculture, we anticipate that this review paper will further encourage and support researchers in their efforts to isolate novel microbial strains, find novel bioindicators, and collaborate with agronomists to modify rooting patterns under various soil management scenarios.

Funding

This research received no external funding.

Data Availability Statement

Not applicable.

Acknowledgments

The authors wish to acknowledge M N Thimme Gowda, Professor and Head Agro-Meteorology, University of Agricultural Sciences, GKVK, Bengaluru-560065 for giving insights in bioirrigation.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Mechanism of bioirrigation through bioindicators in intercropping systems.
Figure 1. Mechanism of bioirrigation through bioindicators in intercropping systems.
Sustainability 15 03542 g001
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Rajanna, G.A.; Suman, A.; Venkatesh, P. Mitigating Drought Stress Effects in Arid and Semi-Arid Agro-Ecosystems through Bioirrigation Strategies—A Review. Sustainability 2023, 15, 3542. https://doi.org/10.3390/su15043542

AMA Style

Rajanna GA, Suman A, Venkatesh P. Mitigating Drought Stress Effects in Arid and Semi-Arid Agro-Ecosystems through Bioirrigation Strategies—A Review. Sustainability. 2023; 15(4):3542. https://doi.org/10.3390/su15043542

Chicago/Turabian Style

Rajanna, Gandhamanagenahalli A., Archna Suman, and Paramesha Venkatesh. 2023. "Mitigating Drought Stress Effects in Arid and Semi-Arid Agro-Ecosystems through Bioirrigation Strategies—A Review" Sustainability 15, no. 4: 3542. https://doi.org/10.3390/su15043542

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