**2. Legume Lectins as a Factor in an Effective Symbiosis**

The full establishment and functioning of the legume-rhizobial symbiosis depends signifi‐ cantly on a number of abiotic, biotic and anthropogenic factors. In particular, the effective‐ ness of symbiotic nitrogen-fixing is affected by temperature, aeration, pH level and moisture content of the substrate, the presence of pesticides, the content of nitrogen and other macroand micronutrients. Of further importance is the virulence and activity of root nodule bacte‐ ria. The study of the influence and the establishment of the degree of importance of different factors to the effectiveness of symbiosis, will determine the conditions, and help to develop methods of optimizing the functioning of the symbiotic systems.

Wide dissemination of lectins in a variety of plants and the presence of these proteins in vir‐ tually all organs and tissues of plant organisms demonstrate the importance of their role in life processes. Initially it was assumed that the presence of lectins was a distinctive feature of seeds from the legume family. However, the number of organisms in which lectins have been found is increasing every day and there is reason to believe that all the above men‐ tioned plants as well as algae, lichens and fungi contain lectins [9-11]. Nevertheless, in the present day researchers focus much of their attention on lectins of leguminous plants, the increased study of which leads us to understand the activity of these proteins in relation to the structure of the latter.

Legume lectins are a large group of carbohydrate binding proteins derived mainly from seeds. Lectins contained in the seeds of legumes, are localized in the proteins and make up approximately 10% of soluble protein extract. In recent years it has been found that these proteins are present in other parts of plants, including stems, leaves, bark, roots, and root nodules. Lectins accumulate in the vacuoles of cells and can come to the surface of the plants. They can also be associated with membranes and cell walls [12-15].

The symbiotic and associative systems of plants and diazotrophs are an example of the evo‐ lution of the interaction of living organisms. Their study is particularly relevant with the im‐ plementation of highly productive and environmentally friendly farming. The biological fixation of molecular nitrogen from the air is a process of fixation and assimilation of nitro‐ gen by microorganisms. It is of great practical importance, since the industrial production of chemical nitrogen fertilizer requires significant use of costly energy resources, which by themselves can be harmful for the environment. A comprehensive study of this problem is necessary due to the need to create new and effective biological preparations. The creation and use of biological agents on the basis of nitrogen-fixing microorganisms is the most justi‐ fiable method of increasing the productivity of plants and the quality of their harvest, which allows maintenance of the natural fertility of soils and the ecological balance of the environ‐ ment. Their use makes it possible to regulate the number and activity of beneficial micro‐ flora in the rhizosphere of crops, and to provide plants with nitrogen fixed from the atmosphere. For example, in addressing the shortage of high-grade protein the key role be‐ longs to the soybean. However, the soils on which the crop is grown for the first time [8] usually do not have nodule bacteria compatible with soybean or bacteria number is small

A Comprehensive Survey of International Soybean Research - Genetics, Physiology, Agronomy and Nitrogen

(up to 20 per gram of soil).

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62

the structure of the latter.

**2. Legume Lectins as a Factor in an Effective Symbiosis**

methods of optimizing the functioning of the symbiotic systems.

The full establishment and functioning of the legume-rhizobial symbiosis depends signifi‐ cantly on a number of abiotic, biotic and anthropogenic factors. In particular, the effective‐ ness of symbiotic nitrogen-fixing is affected by temperature, aeration, pH level and moisture content of the substrate, the presence of pesticides, the content of nitrogen and other macroand micronutrients. Of further importance is the virulence and activity of root nodule bacte‐ ria. The study of the influence and the establishment of the degree of importance of different factors to the effectiveness of symbiosis, will determine the conditions, and help to develop

Wide dissemination of lectins in a variety of plants and the presence of these proteins in vir‐ tually all organs and tissues of plant organisms demonstrate the importance of their role in life processes. Initially it was assumed that the presence of lectins was a distinctive feature of seeds from the legume family. However, the number of organisms in which lectins have been found is increasing every day and there is reason to believe that all the above men‐ tioned plants as well as algae, lichens and fungi contain lectins [9-11]. Nevertheless, in the present day researchers focus much of their attention on lectins of leguminous plants, the increased study of which leads us to understand the activity of these proteins in relation to

Legume lectins are a large group of carbohydrate binding proteins derived mainly from seeds. Lectins contained in the seeds of legumes, are localized in the proteins and make up approximately 10% of soluble protein extract. In recent years it has been found that these The basis of the biological activity of lectins is a reversible reaction with carbohydrates, which defines the different types of biological reactions [16] and their physiological signifi‐ cance. Lectins do not have uniform structural characteristics. In legumes, these proteins are generally composed of two or four subunits with a molecular mass of 25 000 - 30 000 and each having one carbohydrate binding site [15, 17]. Lectins can interact with both mono-and oligosaccharides, as well as the remnants of carbohydrates present in the complex organic substances - glycoproteins, polysaccharides, and glycosides. In the early 1980's there was es‐ tablished a carbohydrate specificity for many legume seed lectins. However indisputable physiological carbohydrate ligands for any of these proteins have not been identified [14].

Over the past half-century many hypotheses were developed regarding the role of lectins in the life of legumes, but so far none of them have been fully confirmed. These proteins were detected due to their ability to bind carbohydrates, and most hypotheses about their func‐ tions are based solely on this wonderful property. Since it is known that the plant may con‐ tain a significant number of lectins, which are localized in different tissues and may form a gene duplication, while carrying out a variety of functions, it is believed that any attempt to dedicate a specific role to these proteins is doomed to failure [14].

An important feature of most of the representatives of the family *Fabaceae* (Legumes) is the ability to enter into symbiotic relationships with rhizobia. They belong to the family of *Rhizo‐ biaceae* (bacteria genera *Rhizobium, Bradyrhizobium, Mezorhizobium, Sinorhizobium, Azorhizobi‐ um*), and form special structures on the roots called nodules. Lectins are considered as a component of the molecular and chemical interactions underlying the formation of symbiot‐ ic structures [6, 9, 11]. The symbiosis between rhizobia and legumes is based on a complex sequence of morphophysiological changes in the cells of both partners. Stages in the develop‐ ment of symbiosis are: preinfection (chemotaxis of bacteria, the exchange of signals, the ad‐ sorption of microsymbionts on the root surface), infection of the roots and development of nodules (penetration, formation of infection threads, the transformation of rhizobia into bac‐ teroids), as well as the functioning of the nodules of nitrogen fixation [9, 18, 19].

Plants throughout their lifespan release various matter into the environment. The presence of lectins has been found in the soil [20], which are secreted during the germination of seeds together with other biologically active substances [21]. It is believed that in the first stages of interaction between rhizobia and legumes an important role is played by the chemotactic re‐ sponse to bacteria. Nodule bacteria and other soil microorganisms, in response to plant exu‐ dates, stimulate the reproduction and active movement of bacteria towards the roots, colonizing the rhizosphere and rhizoplane. A positive chemotaxis of rhizobia on the root exudates and germinating seeds can be either nonspecific (due to the excretion of organic acids, carbohydrates, vitamins), or specific (induced by flavonoids and lectins) [9, 18, 22].

The initial stage of many plant-microbe interactions is recognition of partners, which is largely due to the exchange of molecular signals [23, 24]. This process begins with the exu‐

dation by the host plant of chemicals such as flavonoids and betaines that induce the gene expression of nodulation in rhizobia [25]. Flavonoids induce the transition of bacteria from free-living to those in a symbiotic state [26]. The first chemically-characterized inducer of the expression of nod-gene was luteolin, drawn from the extract of lucern seeds [27]. For each type of rhizobia, plants produce individual flavonoid signals that can stimulate or inhibit the nod-genes of rhizobia [28]. The identification of factors by microorganisms which are re‐ leased by plants initiates physiological processes required for the infection of the host plant. In turn, the microbial signals induce the plant to express the genes required for the forma‐ tion of responses [5, 18, 25, 29]. Amongst the signals produced by the bacteria, the most studied are the lipoсhitooligosaссharide Nod-factors [14, 24]. During the formation of sym‐ biotic relationships lectins on the plant recognize the nod-factor signal released by the bacte‐ ria. It is believed that lectins binding to the bacteria whith the surface of root hairs are able to identify the rhizobial signaling molecules, through surface polysaccharides on the rhizo‐ bia [14, 30]. It has been shown that the lectin of dolichos roots, which is localized on the sur‐ face of root hairs, is able to connect with some of the Nod-factors [14]. Furthermore, from the soybean root tissue there has been found and characterized a chitin-binding protein of the plasma membranes of cells. The presence of this protein and the specificity of the induc‐ tion of the biological response to the binding of ligand *Bradyrhizobium japonicum* indicates its importance in the initiation of response to the binding of chitin in soybean [31].

Therefore Nod-factors play the role of a trigger mechanism for initiating bacterial invasion and nodulation of the plant [24]. Lipopolysaccharides inform the plant about the transition to a symbiotic interaction and the formation of the so called functional areas [32]. Legume plants are unique in their ability to form a response to Nod-factors, which consist of a prepa‐ ration for the penetration of bacteria into the roots. Unfortunately, we do not know how the representatives of this family developed the ability to recognize such signals and how they improved upon these mechanisms [26].

An important stage of preinfection is the adsorption of bacterial cells on the surface of root hairs [33, 34]. The process of attachment of the bacteria from the *Rhizobiaceae* family consists of several stages with the participation of bacterial surface proteins in the first stage (rickad‐ hesin, porin) and polysaccharide fibrils in the second stage. The first stage, involving pro‐ teins, is more decisive for the success of infection than the second stage, mediated by specific cellulose fibrils. The latter promote the retention of bacterial cells on the surface of the plant, but are not necessary for the infection [35].

The adsorption of rhizobia has also been associated with the ability of legumes to synthesize specific glycoproteins called lectins, which bind to polysaccharides on the surface of rhizo‐ bia cells [36, 37]. It is believed [33, 34] that the ability of rhizobia to be adsorbed on the sur‐ face of the roots of the host plant (like the chemotaxis) may be either specific or nonspecific. The initial binding of rhizobial cells to the surface of root hairs occurs by the means of exo‐ polysaccharides (EPS). The degree of affinity between EPS microsymbionts and host plant lectin determines the degree of homology between the symbiotic partners and provides an advantage over other homologous strains in the process of plant infection.

Dazo and his co-writer [38] proposed a model of the attachment of rhizobia to the surface of the roots of dicotyledonous plants. The first phase is non-specific attachment which is char‐ acterized by the fact that the polyvalent host plant lectin binds carbohydrate receptors on the cell surface of nodule bacteria. This results in an intercellular "bridge" between lectin and polysaccharide. The second phase is the "anchoring" of bacteria on the surface of plant cells. This is the phase of specific attachment. The bacterial cells attach to the plant cell, which is the signal for further stages of infection. At this point microfibrils are formed be‐ tween the contacting surfaces which consist of cellulose [35].

dation by the host plant of chemicals such as flavonoids and betaines that induce the gene expression of nodulation in rhizobia [25]. Flavonoids induce the transition of bacteria from free-living to those in a symbiotic state [26]. The first chemically-characterized inducer of the expression of nod-gene was luteolin, drawn from the extract of lucern seeds [27]. For each type of rhizobia, plants produce individual flavonoid signals that can stimulate or inhibit the nod-genes of rhizobia [28]. The identification of factors by microorganisms which are re‐ leased by plants initiates physiological processes required for the infection of the host plant. In turn, the microbial signals induce the plant to express the genes required for the forma‐ tion of responses [5, 18, 25, 29]. Amongst the signals produced by the bacteria, the most studied are the lipoсhitooligosaссharide Nod-factors [14, 24]. During the formation of sym‐ biotic relationships lectins on the plant recognize the nod-factor signal released by the bacte‐ ria. It is believed that lectins binding to the bacteria whith the surface of root hairs are able to identify the rhizobial signaling molecules, through surface polysaccharides on the rhizo‐ bia [14, 30]. It has been shown that the lectin of dolichos roots, which is localized on the sur‐ face of root hairs, is able to connect with some of the Nod-factors [14]. Furthermore, from the soybean root tissue there has been found and characterized a chitin-binding protein of the plasma membranes of cells. The presence of this protein and the specificity of the induc‐ tion of the biological response to the binding of ligand *Bradyrhizobium japonicum* indicates its

A Comprehensive Survey of International Soybean Research - Genetics, Physiology, Agronomy and Nitrogen

importance in the initiation of response to the binding of chitin in soybean [31].

improved upon these mechanisms [26].

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64

the plant, but are not necessary for the infection [35].

Therefore Nod-factors play the role of a trigger mechanism for initiating bacterial invasion and nodulation of the plant [24]. Lipopolysaccharides inform the plant about the transition to a symbiotic interaction and the formation of the so called functional areas [32]. Legume plants are unique in their ability to form a response to Nod-factors, which consist of a prepa‐ ration for the penetration of bacteria into the roots. Unfortunately, we do not know how the representatives of this family developed the ability to recognize such signals and how they

An important stage of preinfection is the adsorption of bacterial cells on the surface of root hairs [33, 34]. The process of attachment of the bacteria from the *Rhizobiaceae* family consists of several stages with the participation of bacterial surface proteins in the first stage (rickad‐ hesin, porin) and polysaccharide fibrils in the second stage. The first stage, involving pro‐ teins, is more decisive for the success of infection than the second stage, mediated by specific cellulose fibrils. The latter promote the retention of bacterial cells on the surface of

The adsorption of rhizobia has also been associated with the ability of legumes to synthesize specific glycoproteins called lectins, which bind to polysaccharides on the surface of rhizo‐ bia cells [36, 37]. It is believed [33, 34] that the ability of rhizobia to be adsorbed on the sur‐ face of the roots of the host plant (like the chemotaxis) may be either specific or nonspecific. The initial binding of rhizobial cells to the surface of root hairs occurs by the means of exo‐ polysaccharides (EPS). The degree of affinity between EPS microsymbionts and host plant lectin determines the degree of homology between the symbiotic partners and provides an

advantage over other homologous strains in the process of plant infection.

The "Lectin" hypothesis which explains the specificity for formation of nodules as resulting from the complementary interaction of the surface structures of bacterial and plant cells. This forms an "antigen-antibody" complex, which was first recognized in the early 1980's. Although initially a controversial subject, Diaz and co-writer [39] provided strong genetic evidence that the root lectins of legumes determine the specifics of symbiosis development. They injected special strains of *Agrobacterium* into the roots of clover through transfer of the psl gene (codes for synthesis of pea lectin). This resulted in clover acquiring the ability to form nodules with pea rhizobia. A similar study ten years later had the same result [40]. Transgenic lucerne plants carrying genes encoding soybean or pea lectin generated a struc‐ ture similar to that of the root nodule in response to incoulation with pea and soybean nod‐ ule bacteria producing the specific Nod-factor. However not all of the structures created were able to form complete infection threads. The results confirm the importance of the presence of lectin in establishing contact between the symbionts and demonstrates the unique role of exopolysaccharides (EPS) in the formation of nodules. *Galega officinalis* and *orientalis* are other examples demonstrating the importance of lectins in legume-rhizobial symbiosis. The validity of this result is confirmed by sequencing the amplifiers of DNA ex‐ tracted from the seeds of the mentioned crops from different geographical locations [41].

Therefore numerous experimental data confirms the important role of lectins in the early stages of the symbiosis and suggests the involvement of these proteins in a variety of phys‐ iological processes of plants.

Once they have prepared each other with molecular signals, the partners begin to form the structural basis of symbiosis - a nodule. This leads to a morphophysiological differentiation of bacterial and plant cells [18, 25]. Nod-factors control the phenotypic changes that occur in the roots of the host plant (the initial stage of symbiosis.) In addition there is also the defor‐ mation and proliferation of root hairs [25], the expression of early nodule genes [5], the in‐ duction of mitotic divisions in the cortex, followed by the onset of the first stages of histogenesis of nodules [42, 43].

Based on the assumption regarding the participation of lectins in various physiological proc‐ esses in the formation of symbiosis, much attention was paid by researchers to the study of the direct effects of plant lectins on the manifestation of the symbiotic properties of nodule bacteria. In particular, it has been shown that the incubation of nodule bacteria *Bradyrhizobi‐ um japonicum* whith soybean root exudates and lectin from its seeds increased the activity of nodule formation in the mutated soybean HS 111, characterized by delayed nodulation, and in doubling the number of nodules formed by wild strain USDA 110 [44]. Later it was

shown that treatment of rhizobia by specific plant lectin increases their virulence and com‐ petitiveness [34], and also increases the quantity of infectious threads in the roots of legumi‐ nous plants [45]. The particular modulating effect of the plant lectin on the formation and functioning of symbiosis has been established, which is manifested in the stimulating effect on partners, homologous data of the proteins, and the neutral or suppressive reaction of lec‐ tin which is not in accordance with the symbionts [46]. Moreover it is shown that the incuba‐ tion of nodule bacteria with homologous lectin has a positive effect on the symbiotic properties of the active strain *Bradyrhizobium japonicum* 634b, and the intensity of the basic physiological processes in plants - nitrogen fixation and photosynthesis. The introduction of the same protein into an inactive strain of rhizobia suspension *Bradyrhizobium japonicum* 604k has a suppressive effect on the symbiotic system which it took part in forming as well as the stated processes [47]. It has also been found that the nature of the influence of homol‐ ogous lectin on the growth of nodule bacteria in a axenic culture and the biosynthesis of ex‐ tracellular carbohydrates [48], the ability of rhizobia to form nodules and their nitrogenase activity, as well as its impact on the productivity of host plants depends on the concentra‐ tion of this protein in the bacterial suspension [49].

Co-incubation of nodule bacteria with lectin enhances soybean growth processes in the early stages of ontogeny as well as in the functioning of the symbiotic system, increases the nitro‐ gen-fixing activity and, consequently, the productivity of plants [47, 49, 50]. This is why in recent years work was done on the selection of the optimal concentration of the homologous lectin, and the length of time of its co-incubation with the culture of *Bradyrhizobium japoni‐ cum* in the production of bacterial fertilizers both in liquid and solid form. Lectin is used as a growth stimulating biologically active substance, which is introduced into the culture of bacteria under certain conditions [51].

In the bacterial model *Bacillus subtilis* where antibiotics were used as metabolic inhibitors, which block the processes of replication, transcription or transmission, there was established an ability of carbohydrate-binding proteins, lectins of plant origin, to have various effects on intracellular processes. Among the presumed processes affected by lectins were reparative functions [52]. It was also found that lectins from the seeds of leguminous plants that have a high molecular weight (> 100 000 Da) can stimulate the respiration of some *Rhizobium*. Ac‐ cording to the authors [53], this effect is induced by the lectins, and significantly increases the interaction of lectin-*Rhizobium* due to the physiological properties of the bacteria.

The presented data gives cause to consider the homologous lectin, not only as a receptor or a signaling molecule in the initial stages of symbiosis, but also as a molecular signal that changes the metabolism of rhizobia, which significantly affects their symbiotic properties and the physiological status of the host plant.

In the process of development of nodules in legume plants, lectins are localized at least in three different places. Their possible functions are related to areas of infection on the surface of the roots. In the nodule primordia lectins can stimulate mitotic activity reducing the threshold of sensitivity to the rhizobial Nod-factors. At the same time, in the central part of the mature nodule, lectins can function as spare nitrogen compounds [54].

It is thought that legume lectins are involved in the formation of nodules. The activating ef‐ fect of lectin on the synthesis of extracellular and capsular polysaccharides of rhizobia is shown, which in turn induces the formation of infection threads [55]. Data for the study of lectin gene expression during the development and functioning of root nodules of lucerne provided evidence supporting the involvement of these proteins in the early stages of the ontogenesis of nodules [56]. A study has been made of a number of genes and plant proteins encoded by them, which play a role in the formation and functioning of root nodules. Among them is a lectin related to the pea *ps*1 gene with the presumed function as a mitotic stimulator and soybean lectin *le*1, which takes part in the attachment of cells [19].

shown that treatment of rhizobia by specific plant lectin increases their virulence and com‐ petitiveness [34], and also increases the quantity of infectious threads in the roots of legumi‐ nous plants [45]. The particular modulating effect of the plant lectin on the formation and functioning of symbiosis has been established, which is manifested in the stimulating effect on partners, homologous data of the proteins, and the neutral or suppressive reaction of lec‐ tin which is not in accordance with the symbionts [46]. Moreover it is shown that the incuba‐ tion of nodule bacteria with homologous lectin has a positive effect on the symbiotic properties of the active strain *Bradyrhizobium japonicum* 634b, and the intensity of the basic physiological processes in plants - nitrogen fixation and photosynthesis. The introduction of the same protein into an inactive strain of rhizobia suspension *Bradyrhizobium japonicum* 604k has a suppressive effect on the symbiotic system which it took part in forming as well as the stated processes [47]. It has also been found that the nature of the influence of homol‐ ogous lectin on the growth of nodule bacteria in a axenic culture and the biosynthesis of ex‐ tracellular carbohydrates [48], the ability of rhizobia to form nodules and their nitrogenase activity, as well as its impact on the productivity of host plants depends on the concentra‐

A Comprehensive Survey of International Soybean Research - Genetics, Physiology, Agronomy and Nitrogen

Co-incubation of nodule bacteria with lectin enhances soybean growth processes in the early stages of ontogeny as well as in the functioning of the symbiotic system, increases the nitro‐ gen-fixing activity and, consequently, the productivity of plants [47, 49, 50]. This is why in recent years work was done on the selection of the optimal concentration of the homologous lectin, and the length of time of its co-incubation with the culture of *Bradyrhizobium japoni‐ cum* in the production of bacterial fertilizers both in liquid and solid form. Lectin is used as a growth stimulating biologically active substance, which is introduced into the culture of

In the bacterial model *Bacillus subtilis* where antibiotics were used as metabolic inhibitors, which block the processes of replication, transcription or transmission, there was established an ability of carbohydrate-binding proteins, lectins of plant origin, to have various effects on intracellular processes. Among the presumed processes affected by lectins were reparative functions [52]. It was also found that lectins from the seeds of leguminous plants that have a high molecular weight (> 100 000 Da) can stimulate the respiration of some *Rhizobium*. Ac‐ cording to the authors [53], this effect is induced by the lectins, and significantly increases

the interaction of lectin-*Rhizobium* due to the physiological properties of the bacteria.

the mature nodule, lectins can function as spare nitrogen compounds [54].

The presented data gives cause to consider the homologous lectin, not only as a receptor or a signaling molecule in the initial stages of symbiosis, but also as a molecular signal that changes the metabolism of rhizobia, which significantly affects their symbiotic properties

In the process of development of nodules in legume plants, lectins are localized at least in three different places. Their possible functions are related to areas of infection on the surface of the roots. In the nodule primordia lectins can stimulate mitotic activity reducing the threshold of sensitivity to the rhizobial Nod-factors. At the same time, in the central part of

tion of this protein in the bacterial suspension [49].

bacteria under certain conditions [51].

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66

and the physiological status of the host plant.

It is established that the proteins extracted from lupine and soybean nodules are capable of hemagglutinating activity. A comparative analysis of the nitrogen-fixation of soybean nod‐ ules and the lectin activity of the protein extracts from these nodules revealed a link be‐ tween these rates in ontogenesis [57]. This data suggests that the proteins which are capable of lectin activity may be directly involved in the processes of nodule functioning.

A hypothesis was made by Antoniuk and his co-writer that the lectin of wheat germs acts as a signal for *Azospirillum brasilense*, which changes the metabolism of the bacteria in a direc‐ tion favorable to the growth and development of the host plant [58, 59]. According to the authors, the level of lectin in the plant depends on a number of conditions and is one of the factors responsible for the variability in the results of wheat inoculation.

Nodule bacteria penetrate the plant cell (as opposed to *Azospirillum*) and transform into bac‐ teroids, which do not divide, but only increase in size. Nonetheless a number of analogies can be made between the influence of the wheat germ lectin on *Azospirillum* and the influ‐ ence of a specific legume lectin on *Rhizobium*. In the first as well as in the second case, the homologous lectin has a positive effect on the symbiotic properties of bacteria, which pro‐ motes a more efficient interaction between the partners in the different symbiotic systems that they create. There remains the little-studied question regarding the participation of lec‐ tins in the functioning of the symbiotic apparatus of legumes, but it is possible that the pres‐ ence of proteins with lectin activity in the nodules is associated with the biosynthesis of proteins (including nitrogenase) in bacteroids and in ensuring conditions for the effective functioning of their symbiotic system.

Legume lectins may influence the receptor and signaling molecules during the stages of the symbiosis. Hemagglutinating activity of the proteins contained in the nodules, varies de‐ pending on the efficiency of their functioning. Specific lectins are able to modify the symbi‐ otic properties of the nodule bacteria which positively affect the physiological status of the host plant and, ultimately, the effectiveness of the symbiotic system. Uncovering the role that lectins have in the functioning of the symbiotic apparatus of legumes requires further research. At the same time these proteins can be regarded as one of the factors for the effec‐ tive symbiosis, which must be considered when developing and implementing new ap‐ proaches to the management of production process in legume plants.
