Selenium uptake, translocation, subcellular distribution and speciation in winter wheat in responses to phosphorus application combined with three kinds of selenium fertilizer

doi:10.21203/rs.3.rs-2331725/v1

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Selenium uptake, translocation, subcellular distribution and speciation in winter wheat in responses to phosphorus application combined with three kinds of selenium fertilizer

https://doi.org/10.21203/rs.3.rs-2331725/v1

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published 27 Apr, 2023

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Background

Selenium (Se) deficiency caused a series of health disorders in human beings, and Se concentration in the edible parts of crops can be improved by altering exogenous Se species. However, the uptake, transport, subcellular distribution and metabolism of selenite, selenate and SeMet influenced by phosphorus (P) has not been well characterized.

Results

The resulted showed that increasing P supply enhanced photosynthesis and then increase the dry matter weight of shoots at selenite and SeMet, and appropriate P combined with selenite increased the dry matter weight of roots by enhancing root growth. At selenite, increasing P supply significantly decreased the concentration and accumulation of Se in roots and shoots. P₁ decreased the Se migration coefficient, which could be attributed to the inhibited distribution of Se in root cell wall but increased distribution of Se in root soluble fraction, as well as the promoted proportion of SeMet and MeSeCys in roots. At selenate, P_0.1 and P₁ significantly increased the Se concentration and distribution in shoots and Se migration coefficient, which could be attributed to the enhanced proportion of Se(IV) in roots but decreased proportion of SeMet in roots. At SeMet, increasing P supply significantly decreased Se concentration in shoots and roots, but increased the proportion of SeCys in roots.

Conclusion

Compared with selenite or SeMet, appropriate P combined with selenite could promote plant growth, reduce Se uptake, alter Se subcellular distribution and speciation, and then affect the Se bioavailability in wheat.

Winter Wheat

Phosphorus

Selenium

Uptake

Translocation

Subcellular distribution

Se species

Selenium (Se) is an essential micro-element for humans and animals, and plays important roles in oxidative stress-related conditions, immune system support, and disease prevention [1, 2]. In recent decades, hidden hunger caused by Se deficiency is commonly found in worldwide [3], including muscle syndrome, Keshan disease, liver disease, cognitive impairment and many cancers [4, 5]. While, excessive Se intake is also harmful, resulting in the health disorders such as hair loss, nervous system disorders and paralysis [6]. The World Health Organization suggest that daily intake of Se for adults should be commended allowance (50–55µg). However, it has been estimated that Se intake amount of about 0.7 billion people in world was lower than the commended value [7]. Studies also have shown that 72% of soil was Se deficient in China [8]. Therefore, Se-enriched fertilizers were applied to improve Se contents in edible plant parts and increase Se intake in human living in Se-deficient areas [9–11].

It is well known that Se availability was influenced by pH, Eh, as well as Se species in soil [12, 13]. Four oxidation states of Se (-II, 0, IV and VI) exist in soil [14], in which, selenide (-II) and Se (0) are difficultly absorbed by roots of plant, selenite (IV) is the main form of Se in acid and neutral soils (pE + pH < 7.5) but selenate (VI) mainly exists in oxidized and alkaline conditions (pE + pH > 15). Plants could uptake both of selenite and selenate, most of which is converted into SeCys and SeMet [15]. Comparing to selenite and selenate, organic Se was safety and beneficial to be uptake by plants [16–18]. Selenomethionine (SeMet) can be absorbed by plants through amino acid transporters [17]. The translocation and metabolism of selenate, selenite and organic Se showed differences after absorption in plants [19, 20]. Further, different forms of Se sources have influenced on Se concentration, species and distribution [21]. For example, selenite was mainly accumulated in roots, but selenate was in shoots [11, 22, 23]. SeMet is the dominant organic Se in plants, and it was translocated from roots to shoots via NRT1.1B in plants [24, 25]. Organic Se could be absorbed into phloem and then translocated to grains via stem, while most of inorganic Se is translocated through xylem to grains [26].

It has been reported that selenite was absorbed by root via passive diffusion or phosphate transporter [11, 23]. Studies demonstrated that translocation of selenite between different organs of plant was promoted by the phosphorus (P) application [27]. P fertilizer increased Se concentration in the soil, and then promoted Se absorption and accumulation in rice [28]. However, several studies have shown that there is an antagonistic effect between phosphate and selenite. Therefore, the interaction of phosphate and selenite is still needed to be verified. Generally, phosphate has little effect on the uptake of selenate due to their chemical dissimilarities. P starvation had no effect on selenate uptake, but decreased selenate concentration in the xylem sap of selenate-treated plants in hydroponic experiments [11]. Some studies reported that P application increased the utilization of selenate, and then increased the Se contents in plants [29, 30]. Compared with inorganic Se, few study was focused on the uptake of organic Se influenced by P application in plants.

Generally, inorganic Se (selenite and selenate) and selenoamino acids (SeMet, SeCys and MeSeCys) are present in plants [13, 22, 31]. Compared with inorganic Se, organic Se is more safety to human health. Studies pointed that different Se sources, P nutrient states and crop variety has significant influence on Se species in plant [13, 20, 32, 33]. Selenite can be quickly assimilated in root, and then converted to organic Se (SeMet, SeCys or SeOMet), however, selenate can was detected in shoot due to quickly mobile in xylem transport [11]. Studies demonstrated P deficiency increased the proportion of MeSeCys in rice, but MeSeCys was not detected when additional P to nutrient solution [13]. The most abundant organic Se is SeMet in wheat and rice, while the main Se species in Indian mustard root is DMeSe and SeMeCys. However, the SeCys and SeMet combined with the protein fraction, affecting the normal structure of protein, and lead to toxicity [20, 34]. Therefore, the proportion of Se species and subcellular fractions in plant is required to assess Se biofortification.

The objectives of this hydroponic study were to investigate (1) the interactive effects of P combined with three kinds of Se fertilizer on the absorption, transport and distribution of Se in wheat; (2) the subcellular fraction and speciation of Se in response to P combined with Se application. Our findings will improve our understanding of the interaction of P and Se in plants and our ability to more effectively regulate the nutritional quality of winter wheat grain via the application of P and Se fertilizers in agricultural practice.

Dry matter weights

Se, P application and their interaction had significant effects on the dry matter weight in shoot and root of winter wheat (P < 0.01; Table S1).

Compared with P_0.01, P_0.1 and P₁ significantly increased the dry matter weights of shoots at selenite and SeMet treatment; there was no significant difference in shoots dry matter weights between different P application rates at selenite treatment (Fig. 1A). Compared with P_0.01, P₁ and P_0.1 significantly decreased the dry matter weights of roots at selenite and SeMet treatment, respectively; both of P_0.1 and P₁ significantly decreased the dry matter weights of roots at selenate treatment (Fig. 1B).

As P was supplied at 0.01 mmol L^− 1, the dry matter weight of shoots in the treatment of selenate and SeMet was significantly higher than that in selenite treatment; at P_0.1, the shoot dry matter weight showed selenite > SeMet > selenite treatment; at P₁, the shoot dry matter weight in selenite and SeMet treatment was higher than that in selenate treatment (Fig. 1A). Compared with selenite and SeMet treatment, selenate application resulted in the highest dry matter weight of roots of wheat at 0.01 mmol L^− 1 P (Fig. 1B); at P_0.1, the dry matter weight of roots in the selenite treatment was higher than that in the selenate and SeMet treatment; at P₁, there was no significant difference in dry matter weight of roots under three kinds of Se fertilizers.

The highest value of dry matter weight in shoot and root was obtained in the treatment of P_0.1-selenite combination and P_0.01-selenate combination, respectively.

Root Morphology

Se, P application and their interaction had significant effects on root length, root surface, root volume, root tip number and root forks, except for the average root diameter influence by P application (P < 0.01 or P < 0.05; Table S2).

At selenite treatment, the total root length, root surface area, root tip number and fork in P_0.1 was higher than both of P_0.01 and P₁ (Fig. 2A, B, E and F). In selenite, compared with P_0.01, P_0.1 and P₁ significantly decreased the root volume and average diameter (Fig. 2C-D). The root length, root surface area, root volume, root surface area, root tip number and fork were significantly decreased with the increasing P application rates at selenate treatment, but the opposite result was observed for the average root diameter (Fig. 2). In SeMet, the total root length, root surface area, root volume, average diameter and fork were firstly increased and then decreased with the increasing P supply, but P_0.1 and P₁ significantly increased the root tip numbers (Fig. 2).

At P_0.01, the total root length, root surface area, root tip number and forks in the selenate treatment were higher than those in the treatment of selenite and SeMet (Fig. 2A, B, E and F); but the opposite result was observed for the average root diameter (Fig. 2D). The root volume in the selenate treatment was higher than those in the treatment of SeMet (Fig. 2C). At P_0.1 and P₁, the total root length, root tip number and forks in the selenite treatment were higher than those in the treatment of selenate and SeMet, but root volume and average diameter in the SeMet treatment were higher than those in the treatment of selenite and selenate (Fig. 2A, C, D, E and F). At P_0.1, the root surface area in the selenate treatment were lower than those in the treatment of selenite and SeMet, but the average diameter in the SeMet treatment were lower than those in the treatment of selenite and selenate (Fig. 2B and D).

The highest value of the total root length, root surface area, root tip number and forks were obtained in the treatment of P_0.1-selenite combination, but the highest value of the root volume and average diameter were obtained in the treatment of P_0.1-SeMet combination.

Photosynthetic

There were significant differences in net photosynthetic rate (P_n), stomatal conductance (G_s), intercellular CO₂ concentration (C_i) and transpiration rate (T_r) between Se, P application as well as their interaction (P < 0.01; Table S3).

Compared with the P_0.01, P_n, G_s, C_i and T_r were significantly increased by P_0.1 and P₁ at each Se fertilizer, with the highest value in P₁ at both treatment of selenite and SeMet (Fig. 3).

At P_0.01, G_s and T_r in selenate were significantly higher than those in selenite and SeMet, and Ci in selenite and SeMet was significantly higher than that in selenite (Fig. 3B-D). At P_0.1 and P₁, G_s, C_i and T_r in SeMet were significantly higher than those in selenite and selenite, but P_n in selenite was significantly higher than that in selenite and SeMet at P_0.1.

The highest value of the P_n was obtained in the treatment of P_0.1-selenite combination, but the highest value of the G_s, C_i and T_r were obtained in the treatment of P_0.1-SeMet combination.

P Concentration And Accumulation In Wheat

Se, P application as well as their interaction had significant effects on P concentration and accumulation in shoots and roots, except for the P concentration and accumulation in root influence by Se fertilizers (P < 0.01; Table S4).

Compared with P_0.01, P_0.1 and P₁ significantly increased P concentration and accumulation in shoots and roots at selenite and SeMet (Fig. 4). At selenate, P_0.1 and P₁ significantly increased P concentration and accumulation in shoots but decreased P accumulation in roots comparing to P_0.01.

At P_0.01, the roots P concentration and accumulation in selenate was higher than that in selenite and SeMet (Fig. 4B and D). At P_0.1, P concentrations of shoots in SeMet and selenate was higher than that in selenite (Fig. 4A). At P₁, P concentrations of shoots in SeMet was higher than that in selenite and selenate, but P concentrations of roots in selenite was higher than that in selenate and SeMet (Fig. 4A and B). At P_0.1 and P₁, the shoot P accumulation in SeMet was higher than that in selenite and selenate, but the highest value for the roots P accumulation was obtained in selenite (Fig. 4C and D).

The highest value of the shoots P concentration and accumulation were obtained in the treatment of P₁-SeMet combination, but the highest value of the roots P concentration and accumulation were obtained in the treatment of P₁-selenite combination.

Se Concentration And Accumulation In Wheat

Se, P application and their interaction had significant effects on Se concentration and accumulation in shoots and roots (P < 0.01; Table S5).

Compared with P_0.01, P_0.1 and P₁ significantly decreased the shoots Se concentration and roots Se concentration and accumulation at selenite and SeMet as well as shoots Se accumulation at selenite, but increased shoots Se accumulation at SeMet (Fig. 5). Nevertheless, P_0.1 and P₁ significantly increased shoots Se concentration, and P₁ significantly increased roots Se concentration at selenate (Fig. 5A and B).

At each P application rate, Se concentrations and accumulations in shoots and roots in SeMet were significantly higher than those in selenite and selenate (Fig. 5).

The highest value of the Se concentration in shoots and roots and Se accumulation in roots were obtained in the treatment of P_0.01-SeMet combination, but the highest value of the Se accumulation in shoots was obtained in the treatment of P₁-SeMet combination.

P, Se Translocation And Distribution In Wheat

Se, P application and their interaction had significant effects on the P and Se migration coefficient, except for the P migration coefficient influenced by P application (P < 0.01; Table S6).

At selenite, there was no obvious difference in P migration coefficient between different P application rates; but at selenate and SeMet, P_0.1 and P₁ significantly increased P migration coefficient comparing to P_0.01 (Fig. 6A). Comparing with P_0.1, P_0.1 and P₁ had no significant influence on the Se migration coefficient at selenite and SeMet, but significantly increased that at selenate (Fig. 6B). At P_0.01, P migration coefficient in selenate was significantly lower than that in selenite, but at P_0.1 and P₁, P migration coefficient in selenate was significantly higher than that in selenite (Fig. 6A). At each P application rate, the Se migration coefficient in was significantly higher than that in selenite and SeMet (Fig. 6B). The highest value of the P migration coefficient was obtained in the treatment of P₁-selenate combination, but the highest value of the Se migration coefficient was obtained in the treatment of P_0.1- selenate combination.

At selenate and SeMet, P_0.1 and P₁ significantly increased the distribution of P in shoots comparing to P_0.01; at selenite, the distribution of P in roots was higher than that in shoot (Fig. 7A). At P_0.01, the distribution of P in shoots in selenite was higher than in selenate and SeMet, but the opposite result was observed at P_0.1 and P₁ (Fig. 7A). At selenite and selenate, the distribution of P in shoots was firstly increased and then decreased with the increasing P supply (Fig. 7B). At each P supply level, the distribution of Se in shoots in the selenate treatment was higher than that in the selenite and SeMet (Fig. 7B).

Se Subcellular Fraction And Distribution In Wheat

Se, P application and their interaction had significant effects on the subcellular fraction of Se in shoot and root (P < 0.01; Table S7).

Compared with P_0.01, P_0.1 and P₁ significantly decreased Se concentrations in each fraction of shoots and roots at selenite; at selenate, there was no pronounced differences in the Se concentrations in each fraction of shoots and roots between different P application rates; at SeMet, Se concentrations in shoots cell organelle and in roots cell wall were significantly decreased by P_0.1 and P₁, and Se concentrations in soluble fraction of shoots and roots were significantly decreased by P₁ (Table 1). The treatment of SeMet had higher Se concentrations in each fraction of shoots and roots than the treatment of selenite and selenate.

In all the treatment, the proportion of Se in cell wall of shoots and roots was higher than that in cell organelle and soluble fractions, except for that in P_0.01-selenite (Fig. 8). At selenite, Se proportion in cell wall and cell organelle of shoots was increased but Se proportion in soluble fractions of shoots was decreased with the increasing P application levels (Fig. 8A); however, P_0.1 and P₁ decreased Se proportion in cell wall of roots but increased Se proportion in soluble fractions of roots comparing to P_0.01 (Fig. 8B). At selenate, Se proportion in cell wall and cell organelle of shoots and roots was decreased but Se proportion in soluble fraction was increased with increasing P supply levels. At SeMet, compared with P_0.01, P₁ increased Se proportion in cell wall of shoots, but decreased Se proportion in soluble fractions of shoots; increasing P supply reduced Se proportion in cell wall of roots but increased Se proportion in cell organelle and soluble fractions of roots. At each P supply level, Se proportion in cell wall of shoots in selenite was higher than that in selenate and SeMet, Se proportion in soluble fraction of shoots in selenate was higher than that in selenite and SeMet, but Se proportion in cell organelle of shoots in SeMet was higher than that in selenite and selenate (Fig. 8A); Se proportion in cell organelle and soluble fraction of roots in selenite were higher than that in selenate and SeMet, but the opposite result was observed for Se proportion in cell wall of roots (Fig. 8B).

Se Species

Se application had significant effects on Se(IV), Se(IV), SeCys, MeSeCys and SeMet concentrations in shoots and roots (P < 0.01; Table S8); P application had significant effects on Se(IV), MeSeCys and SeMet concentrations in shoots as well as Se(IV), SeCys, MeSeCys and SeMet concentration in roots (P < 0.01; Table S8); P and Se interaction had significant effects on Se(IV), Se(IV), MeSeCys and SeMet concentrations in shoots as well as Se(IV), Se(IV), SeCys, MeSeCys and SeMet concentrations in roots (P < 0.01 or P < 0.05; Table S8).

SeCys, MeSeCys and SeMet were detected in shoots and Se (IV), SeCys, MeSeCys and SeMet were detected in roots as Se was supplied as selenite; at selenite, Se (VI), SeCys, MeSeCys and SeMet were detected in shoots and Se(VI), MeSeCys and SeMet were detected in roots; as Se was supplied as SeMet, Se(IV), SeCys, MeSeCys and SeMet were detected in shoots and roots (Table 2).

At selenite, increasing P application rates significantly reduced the concentrations of SeCys, SeMeCys and SeMet in shoots and roots as well as Se (IV) concentrations in roots (Table 2). At selenate, P₁ significantly decreased Se (IV) concentrations of shoots but increased that in roots comparing to P_0.01. At SeMet, P_0.1 and P₁ significantly decreased the concentrations of shoots SeMet, roots MeSeCys and roots SeMet, but increased the roots SeCys concentrations comparing to P_0.01. At each P application level, SeMet treatment had higher concentrations of Se(IV), SeCys, MeSeCys and SeMet in shoots and roots than selenite and selenate treatment, except for the roots SeCys concentrations at P_0.01.

The proportion of SeMet was highest in shoots and roots, except for that in shoots and roots at P₁-selenate (Fig. 9). At selenite, P_0.1 increased SeCys and MeSeCys proportion but decreased SeMet proportion in shoots; the proportion of MeSeCys and SeMet in roots was gradually increased but the SeCys proportion in roots was decreased with increasing P application rates. At selenate, increasing P application levels decreased shoots SeCys and SeMet proportion in shoots and roots, but increased Se (VI) and MeSeCys proportion in each tissue. At SeMet, SeMet proportion in shoots and roots was reduced, but shoots MeSeCys proportion, Se (VI) and SeCys proportion in each tissue was increased with increasing P application level.

Dry matter weight of wheat

In our study, as selenite was supplied, P_0.1 and P₁ significantly increased the shoot dry matter weight comparing to P_0.01, with the highest value in P_0.1 (Fig. 1A). It suggested that appropriate P combined with selenite can promote plant growth, which is consistent with the results of Dai et al. [31] and Kong et al. [35]. However, P_0.1 and P₁ has no significant influence on the dry matter weight of shoots at selenate, but significantly decreased the dry matter weight in roots (Fig. 1A and B). It was inconsistent with the results of [30], who showed that compared with no P fertilizer-treatment, 1 and 2 g·kg^− 1 of P supply significantly increased wheat biomass in the selenate treatment (1 or 2 mg·kg^− 1) in a pot experiment. The reasons for the different results may be related to the differences in Se and P application rates and cultivation methods [22, 36]. At SeMet, increasing P supply also significantly increased the dry matter weight of shoot (Fig. 1A). Our findings suggested that P supply had different effects on wheat growth under different Se fertilizers. Selenite significantly decreased whereas selenate significantly increased the dry matter weight of wheat at P supply treatment [30]. Different Se species as well as their application rates affected crop yield and biomass of plants [33, 37, 38]. In our study, as P was supplied at 0.1 mmol·L^− 1, the dry matter weight in shoots and roots in selenite treatment was higher than that in selenate and SeMet treatment (Fig. 1A and B). This may be due to the fact that appropriate P combined with selenite can promote winter wheat growth and increase grain yield [39, 40]

Root plays a vital role in taking up nutrient elements, increasing the growth and yield of field crops [41]. Plant can adjust their root length, root volume, root surface area, average diameter, root tip numbers and forks to cope with different P conditions [42]. In our study, P_0.1 had higher total root length, root surface, total root volume, average diameter, root tip numbers and forks than other treatment at selenite and SeMet (Fig. 2), which suggested that appropriate P combined with selenite or SeMet promoted the growth of root. Moreover, selenate had higher root dry matter weight as well as root morphological parameters including total root length, root surface, root tip numbers and forks as without P application, but selenite or SeMet had higher root dry matter weight as well as root morphological parameters as P was supplied (Fig. 1B and 2). It indicated that the effects of selenium fertilizer on the root growth were dependent on the P application rates, which was consistent with the results of Boldrin et al. [43]. Low-dose selenite (0.5 and 1 mg·kg^− 1) stimulated plant growth by enhancing root activities [44]. Selenite supply promoted root growth and increased root length, root volume and root effective surface area [45].

Photosynthetic is closely related to plant growth and yield. The net photosynthetic (P_n), stomatal conductance (G_s), intercellular CO₂ concentration (C_i) and transpiration rate (T_r) reflects the plant organic matter accumulation, the size of the stomatal opening, intercellular CO₂ concentration and water evaporation in leaf, respectively. P deficiency decreased ATP synthase activity due to that P is the substrate for ATP synthesis in the chloroplast stroma, and then significantly decreased photosynthetic of barley [46]. In our study, P_0.1 had the highest value of P_n, G_s, C_i and T_r than P_0.01 and P₁ at each Se fertilizer, but there was no significant difference between P_0.1 and P₁ at selenate (Fig. 3). It suggested that medium P application had better effects on photosynthesis, which was consistent with the results of Loudari et al. [47], who pointed out that adequate P significantly increased the photosynthetic parameter and biomass in wheat. There was a closely relationship between the photosynthesis in plants and Se fertilizers [32, 44, 48]. In our study, P_n and G_s in selenite and SeMet was higher than that in selenate as P was supplied at 0.1 mmol·L^− 1 (Fig. 3A-B). Our results suggested that appropriate P combined with selenite or SeMet enhanced photosynthesis further to increase the shoot dry weights.

P And Se Uptake, Translocation And Distribution

P concentration and accumulation of shoot increased with increasing P supply rate at selenite and SeMet (Fig. 4A, C), which was agreement with the results of Li et al.[36]. At selenate, P_0.1 and P₁ significantly increased P concentration and accumulation in shoot, but decreased P accumulation in root, which might be related to the inhibited growth of root by increasing P supply (Fig. 1). Increasing selenate application levels could inhibit P absorption in plant [49]. In our studies, at P₁, the P concentration and accumulation in shoots in SeMet was higher than that in selenite and selenate, but the highest value of roots P concentration and accumulation was obtained in P₁-selenite combination. This result might be attributed to the competition for P transporters (OsPT6) between P and selenite resulted in the more P accumulated in root [13, 23].

The mechanism of uptake and transport of selenite, selenate and SeMet are different in plants [11, 50]. Selenite, selenate and SeMet was absorbed in plants via P treansporters, sulfur (S) transporters and aquaporins [17, 23, 36, 51, 52]. In this study, at the same P supply rate, Se concentration of each wheat organs in SeMet was higher than that in selenite and selenate (Fig. 5), which is agreement with the results of Ali et al. [53], Huang et al.[54] and Eich-Greatorex et al.[55], who pointed out that the uptake rate of SeMet in rice roots was significantly higher than inorganic Se based on the Se concentration-dependent kinetics. Other studies showed that the Se concentration in each tissue of wheat in selenate treatments was higher than that in selenite treatments, which was owe to that selenate had greater bioavailability than selenite [30, 56, 57]. But in our study, Se concentration of shoot and root in selenite treatment was higher than that in selenate (Fig. 5A and B). In contrast, Se concentration of tomato in selenite was ten-fold more than selenate under hydroponic experiment [33]. The current results suggested that the difference in Se concentration between selenite, selenate and SeMet can be explained by their different uptake and metabolisms [11, 58].

Study pointed out that increasing P supply significantly decreased the expression of P transporters (TRIae;Pht1;10 and TRIae;Pht1;11) in roots and further Se concentration in wheat [36]. Our study found that Se concentration in shoot and roots was remarkably decreased with increasing P supply levels at selenite (Fig. 5A, B). It was similar with the results of Wang et al. [13], who found that there was a significantly negative relationship between Se and P concentration in plants. It might be due to that P application inhibited the uptake of selenite or related to the dilution effect by an increase in growth as a result of increased P application [23, 36]. However, P application increased Se concentration in shoot and root at selenate (Fig. 5A, B). It is consistent with the results of Zhang et al. [30], who showed that P fertilizer activated organic matter-bound Se and Fe oxide-bound Se, and then increased soluble Se concentration in soil, which was related to the decreased pH in rich organic matter soil by P application [55]. Thus, the effect of P application on Se concentration and accumulation depended on the type of Se fertilizer [59]. Compared with P_0.01, P_0.1 and P₁ significantly decreased Se concentration and accumulation in shoots and roots at SeMet (Fig. 5). AMF inoculation can promote uptake of SeMet by increasing valid absorption area of roots including total root length, surface area and number of tips [60]. An increase in P supply decreased Se concentration in shoots and roots at SeMet, which might be related to the decreased valid absorption area of roots (the total root length, surface area, volume and forks), as well as the dilution effect resulted by P application [60].

Translocation factor is used to characterize the transfer ability of nutrient elements in plants [61]. In our study, the Se migration coefficient in selenate treatment was significantly higher than that in selenite and SeMet treatment at the same P levels (Fig. 6B), Selenate have a stronger mobility but a little selenite was transported in xylem of plants [62, 63]. Se migration coefficient in selenate treatment was 40–90% higher than that in selenite treatment [53]. Moreover, most Se was distributed in roots in selenite and SeMet treatment, while more Se was found in shoots in selenate treatment (Fig. 7), which was in agreement with the findings of Wang et al. [22]. Previous studies demonstrated that selenite readily accumulates in roots, selenate is more easily transported from root to shoot, and 70% of the total Se exists in straw of rice [43, 64, 65]. The difference in transportation and distribution of selenite, selenate and SeMet in wheat is due to the different metabolism pathways of the three Se forms [66]. In this experiment, compared with P_0.01, P₁ reduced the Se migration coefficient at selenite (Fig. 6B). Migration coefficient of Se was decreased in the higher P concentration plants [57]. Our study found that the migration coefficient and distribution of P and Se in shoots were increased with increasing P supply at selenate (Figs. 6 and 7), which suggested that P and selenate had synergistic effects on the uptake of P and Se in plant. It might be due to the fact that P application enhanced transpiration and further selenate uptake in plants [67].

Subcellular Distribution Of Se

The compartmentalization effect of metals and metal-like elements in cells can greatly affect the level of free heavy metal ion in cells, thus affecting the movement of ion in plants [68, 69]. Se can be considered as both a nutrient and toxic to plants, which is due to that the gap between beneficial and toxic levels is narrow [70, 71]. Roots have a known series of important function in defense caused by toxic element, including binding the elements to cell walls or sequestering them in vacuoles, and then inhibited the translocation of these element to shoots [72, 73]. In our study, the subcellular distribution of Se in each part of wheat was showed by cell wall > cell soluble fraction > cell organelles at three Se fertilizers, regardless of P application (Table 1, and Fig. 8). The cell wall and vacuole (soluble fraction) can sequestrating more metal ion to limit the movement of ion in plants [45]. Previous study demonstrated that cell wall play an important role in binding Se [72]. In addition, Se concentrations in each subcellular fraction were higher in SeMet treatment than in selenite and selenate treatment (Table 1). It might be related to that organic selenium had higher biological activity and absorption efficiency than inorganic selenium [74]. In our study, an increase in P supply decreased the Se concentration in each fraction and the distribution of Se in root cell wall but increased that in root soluble fraction at selenite (Table 1, Fig. 8). These results indicated that increasing the P supply could promote the transmembrane transportation of Se to reduce cell wall bound Se, and then enhanced Se migration coefficient from roots to shoots in wheat [75]. In contrast, an increase P supply level increased the distribution of Se in shoot cell wall and cell organelles and decreased in shoot soluble fraction, which is consistent with the results of [76]. However, P application had no obviously influence on the subcellular distribution of Se at SeMet and selenate (Table 1, and Fig. 8), which might be due to the fact that the uptake pathway of selenate is via sulfate transporters and SeMet is via aquaporins, respectively, but not phosphate transporters [17].

Se Species In Wheat

It is well known that Se is chemically similar to S [77], and Se is converted to organic forms through S metabolic pathway after being absorbed by plant root [78, 79]. Selenate is firstly converted into selenite via two enzymes known as ATP sulfurylase (APS) and APS reductase (APR) in plant [80]; and then selenite is converted into selenide and SeCys by sulfite reductase (SiR) and cysteine synthase; a part of SeCys can be converted into SeMet by a series of enzymes [71, 81]. In our study, the main Se species in wheat was SeMet at selenite and SeMet treatment (Table 2, Fig. 9), which was consistent with the results of previous studies [82, 83]. However, different Se fertilizer had significant effect on Se species in plant. For example, SeMeCys was dominant as selenite was supplied but Se (VI) was dominant with selenate addition in Brassica rapa [84]. SeMet was the major forms in roots and shoots of rice at SeNPs or selenite treatment [85]. Our study showed that selenate (20.8–45.8%) and SeMet (22.5–55.0%) were the main species in roots and shoots of wheat when selenate was applied (Fig. 9), which was consistent with the results of Li et al. [11] and Li et al. [86]. It might be related to that selenate is not completely converted into organic Se as the results of the high uptake rates or storage of Se in vacuoles [20, 87]. Selenite was quickly metabolized to organic Se (SeCys, MeSeCys and SeMet) after absorbing in roots, and then transported to shoots in the form of organic Se [52], which might be the reason for that Se (IV) was only detected in roots (Table 2). At SeMet, SeMet and MeSeCys accounted for more than 79.0% of the total Se, but the low concentrations of Se (IV) and SeCys were detected in roots and shoots (Fig. 9). It was consistent with the results of Wang et al. [88], who showed that SeMet and MeSeCys were the dominant species in organic Se-treated wheat.

In the present study, increasing P supply significantly decreased the concentrations of SeCys, MeSeCys and SeMet in roots and shoots at selenite (Table 2). It might be due to that there was a competitive relationship between SeO₃²⁻ and PO4³⁻ owing to the chemical similarity [23]. SeCys can be converted into MeSeCys via methylation and SeMet by a series of enzymes (i.e., Cystathionine-γ -synthase, Cystathionine-β-lyase, and Met synthase) [39, 81]. In our studies, increasing P supply promoted the transformation of Se species at three kinds of Se fertilizers. It was consistent with the result of Wang et al.[13], who showed that additional P induced a strong reduce in the proportion of SeMet but increased the proportion of Se (VI) and SeCys in roots of rice. It might be related to that P affected the activities of enzymes involving in the Se metabolism, P supply can alter the bioavailability of Se [76]. Moreover, in selenite, an increase of P supply elevated the proportion of SeMet and MeSeCys but decreased that of SeCys in roots at selenite (Fig. 9B). The proportion of Se (VI) was increased, but the proportion of SeMet in shoots and roots was decreased by P supply at selenate (Fig. 9). Previous study suggested that most selenate was distributed in shoots, and little selenate was assimilated into organic forms [11]. At SeMet, increasing P supply decreased the proportion of SeMet, but increased the proportion of SeCys in roots and shoots (Fig. 9). Our study showed that P application had different influence on the proportion of Se species in each organ at three kinds of Se sources, which was probably due to different uptake mechanisms and P supply influenced the Se assimilate through affecting protein synthesis and some enzymes activity [13, 17]. However, further studies on the Se metabolism as mediated by P application are still needed.

The current study demonstrated that the uptake, translocation and distribution of Se in wheat was significantly influenced by P supply under three Se fertilizers. An increase in P supply decreased Se concentration in shoots and roots at selenite and SeMet treatment, while increased Se concentration in shoots and promoted Se translocate form root to shoot at selenate treatment. The percentage of Se in cell wall was higher than that in organelle fraction and soluble fraction under three kinds of Se fertilizer, whereas an increase in P supply inhibited Se distribution in shoot soluble fraction and root cell walls at selenite treatment, and then effected Se migration coefficient. SeMet was the main species at selenite and SeMet treatment, while SeMet and Se(IV) were the main species at selenate treatment, and the highest concentrations of organic Se was found in shoots and roots at SeMet. P supply had different influence on the proportion of Se species in wheat under three kinds of Se fertilizer. Further studies are needed to elucidate the transformation of Se species by P supply.

Plant materials and treatment

A hydroponic experiment was conducted in a greenhouse with controlled environmental condition at about 14/10 h in light/dark regime, 22/18°C in air temperatures, approximate 500 µmol m^− 2 s^− 1 in photon flux density, and 65% in relative humidity.

Winter wheat (Triticum aestivum cv. Bainong 207, purchased from Henan Agricultural High Tech Group Co., Ltd.) seeds were germinated in deionized water at 25℃ for 4 d, after sterilized in 5% (v/v) NaClO solution for 15 min. Then, uniform size wheat seedlings were transplanted into a plastic container containing 2 L nutrient solutions as described previously by Arnon et al. (1940), which was consisted of 945 mg L^− 1 Ca (NO₃)₂·4H₂O, 607 mg L^− 1 KNO₃, 493 mg L^− 1 MgSO₄·7H₂O, 20 mg L^− 1 EDTA-Fe, 2.86 mg L^− 1 H₃BO₃, 1.81 mg L^− 1 MnCl₂·4H₂O, 0.22 mg L^− 1 ZnSO₄·7H₂O, 0.08 mg L^− 1 CuSO₄·5H₂O, and 0.02 mg L^− 1 (NH₄)₆Mo₇O₂₄·4H₂O.

Experiment Design

P was added to the solutions as NaH₂PO₄·2H₂O at three rates: 0.01, 0.1 and 1 mmol L^− 1 and Se was added as Na₂SeO₃, Na₂SeO₄ and SeMet at one rete of 2 µmol L^− 1. Three times was replicated in each treatment. Seedlings were supplied with full-strength nutrient solutions until sampled, except for the quarter-strength and half-strength solutions supplied during the first and second weeks, respectively. The solution was renewed every 3 days to ensure enough nutrients concentration. Solution of 5% HCl was used to dip all vessels for 1 week, and then deionized water was used to wash those vessels more than three times. After cultivation for 21 d, the roots and shoots of 14 seedlings were separated, dried at 70 ± 5°C, and analyzed for dry weights and concentrations of P and Se. The leaves, stems, and roots of other six seedlings were immediately frozen in liquid nitrogen, and then stored at − 20°C for further subcellular fractions and Se speciation analysis.

Analysis Of The Root Morphology

Three 14-d-old seedlings in each treatment were analyzed for root morphological characteristics. According to the method of Nie et al. [89], the root length, root surface area, root volume, average root diameter, root tip number and forks were measured using the root imaging analysis software WinRHI-ZO Version 2009 PRO (Regent Instruments, Quebec City, Canada).

Analysis Of The Photosynthetic Parameter

According to the method of Shi et al. [90], the net photosynthetic (P_n), stomatal conductance (G_s), intercellular CO₂ concentration (C_i) and transpiration rate (T_r) in the leaves of 21-d-old seedlings were measured using the LI-6400XT photosynthetic-fluorescence assay system (USA, LI-COR).

Analysis Of The P Concentration Of Wheat

According to the method of Bao. [91], After being digested with H₂SO₄-HClO₄, vanadate-molybdate-yellow colorimetric method was used to estimate plant phosphorus concentration.

Analysis Of The Se Concentration Of Wheat

According to the method of Liu et al. [28], approximately 0.5 g of plant samples (shoot or root) was digested with HNO₃-HClO₄ (4:1) at 180°C. The digest was then restored to volume with 6 mol L^− 1 HCl, cooled, filtered and brought to volume with deionized water. The Se concentration was determined by hydride generation atomic fluorescence spectrometry (HG-AFS-8220, Beijing Titan Instruments Co., China)

Subcellular Fractions Separation

According to the methods described by Wan et al. [68], approximately 0.4 g of frozen samples (shoot or root) was homogenized in 10 mL extraction buffer containing 1.0 mM dithioerythritol, 250 mM sucrose, and 50 mM Tris-HCl (pH 7.5). The homogenate was centrifuged at 300 g for 10 min, and the residue constituted the cell wall fraction (F1). The supernatant was then centrifuged at 20,000 g for 30 min, and the precipitate and the supernatant formed the organelle fraction (F2) and the soluble fraction (F3), respectively. The soluble fraction was diluted to 50 mL with 5% HNO₃ (GR) prior to elemental determination. All steps were performed at 4◦C.

Se Speciation Determination

According to the methods described by Wang et al. [92], 0.05 g of the samples (shoot or root) was placed 15 mL centrifugal tubes with 5 mL Tris-HCl. After ultrasonic 30 min, 50 mg cellulase and 20 mg protease K were added. The mixture was incubated in an oscillation box with horizontal shaking (250 r·min^− 1) at 50 ℃ ± 2 ℃ for 18 h. after adding 20 mg protease K, the mixture was then incubated in an oscillation box with horizontal shaking (250 r·min^− 1) at 37 ℃ ± 2 ℃ for 18 h. The hydrolysate was centrifuged at 10000 r·min^− 1 for 30min, and the supernatant was filtered through a 0.22 µm cellulose nitrate filter (Millipore, Billerica, MA, United States). Subsequently, the filtrate was stored at -80°C for Se speciation analysis, using high performance liquid chromatography-ultraviolet treatment hydride generation-atomic fluorescence spectrometry (HPLC-UV-HG-AFS; SA-50, Ji tian Instruments Co., Beijing, China).

Data analysis

All data were statistically analyzed by two-way ANOVA with LSD multiple comparison at a 5% level (P < 0.05) using SPSS 18.0.

Acknowledgments

We sincerely thank Henan Agricultural High Tech Group Co., Ltd. for kindly providing seeds of Bainong 207.

Authors’ contributions

HL and ZN conceived and designed the experiments. HP, GL, CL and HS performed the experiments. ZN and CH analyzed the data. CH wrote the paper. All authors have read and approved the submitted manuscript.

Funding

This research was financially supported by the Key Scientific Research Plans of Colleges and Universities in Henan Province (Program No. 19zx007), the National Key R&D Program of China (Program No. 2021YFD1700900) and the Natural Science Foundation of Henan Province (Program No. 222300420464).

Availability of data and materials

The datasets generated or analyzed during the current study are available from the corresponding author on reasonable request.

Ethics approval and consent to participate

Experimental research and field studies on plants (either cultivated or wild), including the collection of plant material, complied with relevant institutional, national, and international guidelines and legislation. The experiments did not involve endangered or protected species.

Consent for publication

Not applicable.

Competing interests

The authors declare that they have no competing interests.

Author details

¹Resources and Environment College, Henan Agricultural University, No. 63, Nongye Road, Jinshui District, Zhengzhou 450002, Henan Province, China.

²Department of Chemistry and Biochemistry, Texas Tech University, Lubbock, TX 79409, USA.

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No competing interests reported.

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Selenium uptake, translocation, subcellular distribution and speciation in winter wheat in responses to phosphorus application combined with three kinds of selenium fertilizer

Status:

Journal Publication

Version 1

Abstract

Background

Results

Conclusion

Figures

Background

Results

Dry matter weights

Root Morphology

Photosynthetic

Se Concentration And Accumulation In Wheat

Se Subcellular Fraction And Distribution In Wheat

Discussion

Dry matter weight of wheat

P And Se Uptake, Translocation And Distribution

Subcellular Distribution Of Se

Se Species In Wheat

Conclusion

Materials And Methods

Plant materials and treatment

Experiment Design

Analysis Of The Root Morphology

Analysis Of The Photosynthetic Parameter

Analysis Of The P Concentration Of Wheat

Analysis Of The Se Concentration Of Wheat

Subcellular Fractions Separation

Se Speciation Determination

Data analysis

Declarations

References

Tables

Additional Declarations

Supplementary Files

Status:

Journal Publication

Version 1