CN108401712B - Application of glutamine in low-phosphorus rhizosphere acidification - Google Patents
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Abstract
The invention relates to application of glutamine in low-phosphorus rhizosphere acidification, belonging to the technical field of agriculture. Glutamine is an important small amino acid molecule, other than that involved in the photorespiration process andbesides nitrogen assimilation, the low phosphorus state in the plant leaves can be sensed as signal molecules and transmitted to roots from the leaves, so that the acidification level of plant roots is adjusted, and the absorption and utilization of inorganic phosphorus in soil by plants are influenced. In addition, mutants associated with glutamine accumulationp444There is also a similar phenotype of rhizosphere acidification deletion, further suggesting that endogenous glutamine is involved in rhizosphere acidification as a signal molecule. Glutamine is applied to the roots of the wheat seedlings, so that the low phosphorus stress resistance can be improved. Functional analysis and identification of glutamine establish reaction characteristics and molecular mechanism of glutamine in plant phosphorus nutrition utilization, and the method has wide application prospect.
Description
Technical Field
The invention relates to agricultural technology, in particular to application of glutamine in low-phosphorus rhizosphere acidification.
Background
Phosphorus, as one of the macronutrient elements, not only participates in the formation of the basic structure of plants, but also participates in the regulation of various physiological processes of plants. Phosphorus required by plant growth and development is mainly absorbed from soil, although the content of total phosphorus in the soil is not low, most of phosphorus in the soil is combined with calcium, iron, aluminum and the like to form insoluble and low-mobility ineffective phosphorus, so that the soluble effective phosphorus which can be absorbed and utilized by plants is very little, and the lack of phosphorus nutrition is a common adversity stress in the plant growth and development process. Plants have developed many strategies to cope with low phosphorus stress during long-term evolution. For example: the low phosphorus can induce the length and the number of the lateral roots and the root hairs of the plants to be increased, and the absorption area of phosphorus nutrition is enlarged; the low phosphorus can also induce the roots of plants to secrete substances such as organic acid, enzymes and the like, so that the insoluble and directly unavailable phosphorus is converted into soluble phosphorus; the low phosphorus can also induce the expression of a plurality of phosphorus starvation response genes, improve the content and the activity of phosphorus transport proteins, and promote the transmembrane transport and absorption of phosphorus, thereby realizing the high-efficiency utilization of the phosphorus nutrition of the soil by plants.
Under the stress of low phosphorus, the arabidopsis rhizosphere is acidified, the pH value of the rhizosphere can be reduced to be below 5.5, and insoluble phosphorus can be converted into soluble phosphorus by low pH, so that the high-efficiency absorption of plants to soil phosphorus nutrition is promoted. The color of the bromocresol purple of the acid-base indicator is changed from yellow to purple between pH5.2 and 6.8, so the acidification process of the rhizosphere can be visually shown by the color change of the bromocresol purple. Through observing the color change of the acidification culture medium containing bromocresol purple, the rhizosphere acidification deletion arabidopsis mutant plant under the low-phosphorus condition can be intuitively and efficiently screened, and a foundation is laid for revealing a regulation and control mechanism for efficient absorption and utilization of phosphorus nutrition on a molecular level.
We found a mutant causing glutamine accumulation by mutant screeningp444The experiments of regulating and controlling the rhizosphere acidification reaction of arabidopsis thaliana, grafting and injecting leaves prove that the glutamine remote signal molecules in the leaves regulate and control the rhizosphere acidification reaction. The root irrigation experiment of the wheat seedling further verifies the function of improving the low-phosphorus stress resistance of the crop. The revealing of the function of the small molecular glutamine has wide application prospect in promoting the absorption of phosphorus nutrition of soil.
Disclosure of Invention
The invention aims to provide application of arabidopsis thaliana small-molecule glutamine in improving the utilization efficiency of soil phosphorus nutrition, and the small molecule participates in mediating and regulating rhizosphere acidification reaction induced by low phosphorus and release of phosphorus elements in rhizosphere soil, so that a chance is provided for realizing efficient utilization of soil phosphorus nutrition.
The invention also aims to provide a novel method for applying the small molecular amino acid-glutamine to the aspect of low phosphorus stress resistance of plants so as to expand the application range of the glutamine and realize high efficiency of soil phosphorus nutrition.
The technical scheme of the invention is realized as follows:
the small molecule amino acid-glutamine, C thereof5H10N2O3And the molecular weight is 146.15. The small molecular amino acid is used as a signal molecule and participates in a rhizosphere acidification process induced by low phosphorus.
Through a pH indicator bromocresol purple in-situ color development method and exogenous application of glutamine and glutamic acid with different concentrations, the inventor finds that the glutamine participates in a low-phosphorus induced rhizosphere acidification process. The petiole injection experiment further proves that glutamine is a small molecule regulation signal for mediating the rhizosphere acidification process from the leaf part.
Liquid chromatography-mass spectrometry experiments show that related mutantsp444Resulting in an increase in glutamine content; mutant accumulating glutaminep444Analysis of the phenotype and rhizosphere pH values of (a) shows that endogenous glutamine is also involved in the low phosphorus induced rhizosphere acidification process.
The grafting experiment shows that the arabidopsis mutantp444And low phosphorus induced rhizosphere acidification is remotely regulated from aerial leaves. The small-molecule glutamine plays a role in remotely regulating and controlling signals in the rhizosphere acidification reaction induced by low phosphorus.
Wheat experiments show that through analysis of signal functions and action mechanisms of glutamine, research basis and premise are provided for obtaining a new transgenic phosphorus efficient crop strain and improving the stress resistance of crops under low phosphorus stress.
The application of the glutamine in the aspect of low-phosphorus rhizosphere acidification comprises the following steps: adding glutamine into water, shaking to obtain glutamine water solution, and root irrigation of crop with glutamine water solution.
The concentration of the glutamine water solution is 50-500 mu M.
The root irrigation method comprises the following specific operations: in hydroponic crop applications, aqueous glutamine solution is added to the culture medium to achieve a final glutamine concentration of 50-500 μ M, completing root irrigation.
The glutamine is used for inhibiting rhizosphere acidification reaction and is injected at the position of a crop leaf stalk to form 6-10 muL glutamine with the concentration of 50-500 muM.
The beneficial effect that this technical scheme can produce:
(1) the application of glutamine on the roots is beneficial to the low-phosphorus stress resistance of plants, and especially has obvious promotion effect on the growth of the roots in the initial stage under the low-phosphorus condition; in terms of mechanism, under the condition of low phosphorus, glutamine is used as a signal molecule and can play a role in remote control in leaves, so that the rhizosphere acidification of plant roots is reduced, and the tolerance of low phosphorus stress is improved.
(2) Phosphorus is used as an essential nutrient element in the growth process of crops, is easy to lack, and can be absorbed by the crops little after being applied by the conventional crop fertilizer, in the case of low phosphorus, the crops have the undesirable characteristics of short and small plants, stiff plants, slow leaf emergence, few and small leaves and the like, in the face of the common crop diseases, the growers often adopt fertilizer spreading operation, and the application of the fertilizer increases the burden of the growers on the one hand and has slow effect on the other hand, and sometimes causes irreversible damage, the technical personnel of the application aim to solve the technical problem and find that the glutamine has great effect on resisting low phosphorus stress, therefore, the harm caused by phosphorus deficiency can be solved by applying glutamine, and the method is convenient, can be sprayed, root-irrigated or injected, practically and efficiently solves the technical problem and brings huge commercial application value.
Drawings
FIG. 1 shows mutantsp444And complementing the rhizosphere acidification phenotype of the transgenic plant.
FIG. 2 shows WT andp444differences in glutamine content in mutants.
FIG. 3 is a graph showing rhizosphere acidification reaction characteristics of glutamic acid and glutamine applied at different concentrations in vitro under low phosphorus stress.
FIG. 4 is the low phosphorus rhizosphere acidification reaction profile of wild-type petiole injected glutamine.
FIG. 5 shows WT andp444rhizosphere acidified phenotype after grafting.
FIG. 6 shows the low phosphorus stress resistant phenotype of wheat seedlings after glutamine application.
Detailed Description
The technical solutions of the present invention will be described clearly and completely with reference to the following embodiments of the present invention, and it should be understood that the described embodiments are only a part of the embodiments of the present invention, and not all of the embodiments. All other embodiments, which can be obtained by a person skilled in the art without inventive effort based on the embodiments of the present invention, are within the scope of the present invention.
Rhizosphere acidification deletion mutantp444The arabidopsis mutant obtained by screening of the inventor shows that through map-based cloning analysis,p444the mutant has a mutation site located atGLU1。The anaplerotic strain is analyzed by a bromocresol purple indicator in-situ color development method, the acidification degree of the root base of the anaplerotic strain is basically consistent with that of a wild type in phenotype and acidification quantification after low phosphorus stress (figure 1), and the result shows thatp444The mutant is composed ofGLU1Caused by the mutation of the gene, and the gene is a mutant,GLU1the gene is involved in regulating and controlling the rhizosphere acidification process induced by low phosphorus.
It has been shown by the research that,GLU1the gene encodes glutamate synthetase, which catalyzes the conversion of glutamine to glutamate,the The mutation of the enzyme may affect the glutamine and glutamic acid content in arabidopsis thaliana. In order to verify whether the gene mutation can cause the content change of endogenous glutamine and glutamic acid of plants, the experiment adopts a liquid chromatography and mass spectrometry combined method to determine WT and the mutantp444Glutamine content at different sites.
Wild type and mutant growing normally for about 10 dp444Seedlings were treated with normal and low phosphorylation. After about 30 h of culture, after the acidification phenotype is obvious, roots and leaves of the wild type and the mutant are respectively collected. The leaf weight was 0.1 g, the root weight was 0.05 g (considering that part of the acid secreted from the root system remained on the root after treatment, it was washed 2 times with double distilled water, placed on absorbent paper and weighed after the surface moisture was sucked dry). Then the material is placed in a 2 mL centrifuge tube, quick frozen by liquid nitrogen, ground by a sample grinder (60 Hz, 90 s), fully ground, dissolved by a mobile phase (1/1000 glacial acetic acid, 1 mL glacial acetic acid is added into 1L ultrapure water, and the pH is between 2 and 3) with a corresponding volume (1 mL is added in leaf treatment and 0.5 mL is added in root treatment), and subjected to 75 ℃ water bath for 15 min. Centrifuging at 12000 rpm for 20 min, and suckingAnd (4) supernatant fluid. Filtering the supernatant with a filter membrane with the diameter of 0.22 μm, collecting, and measuring the content of glutamine by LC-MS. The reaction conditions are as follows: aminex HPX-87H column, flow rate 0.4 mL/min, sample size 10 u L. And manually integrating the sample peaks according to the peak output time and area of the standard sample and the corresponding mole number to perform quantitative analysis.
The results are shown in FIG. 2. The wild type has higher glutamine in leaves than roots under normal and low-phosphorus stress, and the glutamine content in leaves and roots is reduced under low-phosphorus conditions; mutantsp444Under normal conditions, the glutamine content in the leaves is higher than that in the roots, but under low-phosphorus conditions, the glutamine content in the roots is higher than that in the leaves, and under low-phosphorus conditions, the glutamine content in the leaves is obviously reduced, and the glutamine content in the roots is obviously increased. The results show that the method has the advantages of high yield,p444the mutation results in an increase in the endogenous glutamine content of the plant, and also indicates that glutamine has a positive correlation with low-phosphorus induced rhizosphere acidification.
And analyzing the stress of the rhizosphere acidification reaction of arabidopsis thaliana by applying glutamine with different concentrations to the roots by using a bromocresol purple indicator in-situ color development method.
As shown in FIG. 3, glutamic acid and glutamine were added to a medium containing pH developer bromocresol purple at different concentrations (0. mu.M, 5. mu.M, 50. mu.M, 500. mu.M) under low-phosphorus (12.5. mu.M Phosphate (Pi)) treatment conditions, and the effect of glutamic acid and glutamine on the rhizosphere acidification reaction of Arabidopsis thaliana was analyzed. Wild type Arabidopsis thaliana (WT) was cultured on MS solid medium for 10 days, and transferred to low-phosphorous MS medium containing bromocresol purple for 30 hours. Observation and analysis revealed that the application of various concentrations of glutamic acid had no significant effect on rhizospheric acidification under low phosphorus conditions, whereas at the concentrations of glutamine applied of 50 μ M or more, the rhizospheric acidification phenotype of WT was completely absent. Thus, glutamine inhibits rhizosphere acidification reaction of Arabidopsis under low phosphorus conditions.
Based on the above observations, the inventors speculate that glutamine is a small molecule regulatory signal. Therefore, petiole injection experiments were performed to verify this inference. The relevant experimental procedures are briefly described as follows:
wild type seedlings that grew normally for about 10 d were treated in low phosphorous treatment (12.5. mu.M Phosphate (Pi)) medium for about 30 h. After different treatments of the aerial parts, acidification was observed (see FIG. 4). 6 muL of 50 muM glutamine was injected at the petiole. After the treatment, the seedlings are cultured for about 30 h, and then the rhizosphere acidification condition of the seedlings under the condition of low phosphorus is observed. As a result, after 6 muL of 50 muM glutamine was injected at the petiole, rhizosphere acidification reaction was inhibited. We therefore conclude that: the loss of rhizosphere acidification may be the result of glutamine accumulation in the leaf.
The inventor speculates that glutamine is used as a small molecule to remotely control the rhizosphere acidification process, and in order to further confirm that glutamine is a signal molecule for controlling the underground part of the overground part in the low-phosphorus rhizosphere acidification reaction, in order to verify the speculation, the cutting grafting experiment is used:
wild type and mutant growing normally for about 6 dp444Cutting the upper third of seedling hypocotyl, and collecting the aerial parts and underground parts of wild type, the aerial parts of wild type and mutantp444Underground part of、Mutantsp444The above-ground part and the wild-type underground part, mutantp444The overground part and the underground part are grafted, the culture medium used for grafting is 1.0 percent of an inclined MS culture medium (the inclination angle is 5-10 degrees, then the culture medium is vertically placed and is narrow at the top and wide at the bottom), the successfully grafted seedlings are placed on a normal and low-phosphorus chromogenic culture medium and are treated for about 20 hours (as shown in figure 5), and as a result, when the local upper part is a wild type, rhizosphere acidification can occur under the low-phosphorus condition no matter whether the underground part is a wild type or a mutant; when the local part is mutant, rhizosphere acidification does not occur under low-phosphorus conditions no matter whether the underground part is wild type or mutant. It is thus shown that the signal regulating rhizosphere acidification is likely to be from the aerial part, i.e.p444The rhizosphere acidification reaction involved in low phosphorus induction is remotely regulated from the aerial parts of the leaves.
To verify the role of glutamine in crop resistance to low phosphorus stress, wheat seedlings were rooted in glutamylAmine to verify its effect. With 0.1% HgCl2Sterilizing wheat seeds (the variety is Chinese spring). Soaking seeds in distilled water for 24 h, heating the seeds in an incubator at 28 ℃, keeping the humidity at 60%, illuminating for 14 h/dark for 10 h, and accelerating germination for 72 h. The 96-well PCR plates with holes cut through the lower part are respectively placed in a normal culture solution (MS), 1 seedling is planted in each tube, and 20 tubes are used as a whole. And (3) under normal conditions, adding 50 mu M glutamine, treating the seedlings under low phosphorus (12.5 mu M Phosphate (Pi)), treating the seedlings under low phosphorus treatment and adding 50 mu M glutamine, and photographing and counting after growing for 7 d. The results are shown in fig. 6, and it is found that after 50 μ M glutamine is applied under low-phosphorus conditions, the fresh weight, the root length, the number and the length of lateral roots of the seedling roots are all reduced compared with the treatment only with low phosphorus, and a low-phosphorus resistant phenotype is shown, so that the glutamine can improve the low-phosphorus stress resistance of crops and has a wide application prospect.
The above description is only for the purpose of illustrating the preferred embodiments of the present invention and is not to be construed as limiting the invention, and any modifications, equivalents, improvements and the like that fall within the spirit and principle of the present invention are intended to be included therein.
Claims (7)
1. The application of glutamine in low-phosphorus rhizosphere acidification is characterized in that: the glutamine is used as a signal molecule to regulate and control the rhizosphere acidification reaction of the arabidopsis with low phosphorus induction and improve the application of the arabidopsis in the aspect of phosphorus element lack tolerance;
the method comprises the following steps: adding glutamine into water, oscillating and shaking up to obtain a glutamine aqueous solution, and irrigating roots of crops by the glutamine aqueous solution; the concentration of the glutamine water solution is 50-500 mu M.
2. The use of glutamine according to claim 1 for low phosphorus rhizosphere acidification: the glutamine plays a role in remotely regulating and controlling signals in the rhizosphere acidification reaction induced by low phosphorus.
3. The use of glutamine according to claim 2 for low phosphorus rhizosphere acidification: the glutamine is a small molecule regulatory signal that mediates the rhizosphere acidification process from the leaf site.
4. The use of glutamine according to claim 2 for low phosphorus rhizosphere acidification: the glutamine is a signal molecule for regulating the underground part of the overground part in the low-phosphorus rhizosphere acidification reaction.
5. The use of glutamine according to claim 1 for low phosphorus rhizosphere acidification: the use of said glutamine to increase the resistance of crops to low phosphorus stress.
6. The use of glutamine according to claim 1 for low phosphorus rhizosphere acidification: the root irrigation method comprises the following specific operations: in hydroponic crop applications, aqueous glutamine solution is added to the culture medium to achieve a final glutamine concentration of 50-500 μ M, completing root irrigation.
7. The use of glutamine according to claim 1 for low phosphorus rhizosphere acidification: the application of inhibiting rhizosphere acidification reaction is to inject 6-10 muL glutamine with the concentration of 50-500 muM at the leaf stalk of crops.
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| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| GB869948A (en) * | 1959-03-18 | 1961-06-07 | Ajinomoto Kk | Process for producing glutamic acid from fermentation broth |
| CN101407824A (en) * | 2008-11-24 | 2009-04-15 | 昆明理工大学 | Construction and use of plant expression vector of Arabidopsis thaliana cytoplasm type glutamine synthetase gene |
| CN101781660A (en) * | 2009-01-21 | 2010-07-21 | 河南大学 | Application of arabidopsis gene IAR1 in aspects of iron nutrient and drought resistance |
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| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| GB869948A (en) * | 1959-03-18 | 1961-06-07 | Ajinomoto Kk | Process for producing glutamic acid from fermentation broth |
| CN101407824A (en) * | 2008-11-24 | 2009-04-15 | 昆明理工大学 | Construction and use of plant expression vector of Arabidopsis thaliana cytoplasm type glutamine synthetase gene |
| CN101781660A (en) * | 2009-01-21 | 2010-07-21 | 河南大学 | Application of arabidopsis gene IAR1 in aspects of iron nutrient and drought resistance |
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