CN115067335B - Application of chondroitin sulfate as active ingredient in preparation of plant immunity inducer - Google Patents
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- 229920001287 Chondroitin sulfate Polymers 0.000 title claims abstract description 88
- SQDAZGGFXASXDW-UHFFFAOYSA-N 5-bromo-2-(trifluoromethoxy)pyridine Chemical compound FC(F)(F)OC1=CC=C(Br)C=N1 SQDAZGGFXASXDW-UHFFFAOYSA-N 0.000 title claims abstract description 86
- 229940059329 chondroitin sulfate Drugs 0.000 title claims abstract description 86
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Classifications
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
- A01N43/00—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds
- A01N43/02—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one or more oxygen or sulfur atoms as the only ring hetero atoms
- A01N43/04—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one or more oxygen or sulfur atoms as the only ring hetero atoms with one hetero atom
- A01N43/14—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one or more oxygen or sulfur atoms as the only ring hetero atoms with one hetero atom six-membered rings
- A01N43/16—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one or more oxygen or sulfur atoms as the only ring hetero atoms with one hetero atom six-membered rings with oxygen as the ring hetero atom
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01P—BIOCIDAL, PEST REPELLANT, PEST ATTRACTANT OR PLANT GROWTH REGULATORY ACTIVITY OF CHEMICAL COMPOUNDS OR PREPARATIONS
- A01P1/00—Disinfectants; Antimicrobial compounds or mixtures thereof
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01P—BIOCIDAL, PEST REPELLANT, PEST ATTRACTANT OR PLANT GROWTH REGULATORY ACTIVITY OF CHEMICAL COMPOUNDS OR PREPARATIONS
- A01P3/00—Fungicides
Landscapes
- Life Sciences & Earth Sciences (AREA)
- Wood Science & Technology (AREA)
- Pest Control & Pesticides (AREA)
- Plant Pathology (AREA)
- Environmental Sciences (AREA)
- Engineering & Computer Science (AREA)
- Zoology (AREA)
- Chemical Kinetics & Catalysis (AREA)
- Chemical & Material Sciences (AREA)
- General Chemical & Material Sciences (AREA)
- General Health & Medical Sciences (AREA)
- Dentistry (AREA)
- Health & Medical Sciences (AREA)
- Agronomy & Crop Science (AREA)
- Microbiology (AREA)
- Mycology (AREA)
- Agricultural Chemicals And Associated Chemicals (AREA)
Abstract
The invention discloses application of chondroitin sulfate as an active ingredient in preparation of a plant immunity inducer, belongs to the technical field of agricultural biopesticides, and provides novel application of Chondroitin Sulfate (CS) in inducing plant disease resistance, wherein the chondroitin sulfate can induce accumulation of plant endogenous salicylic acid; exciting active oxygen burst; improving the resistance of plants to pathogenic bacteria.
Description
Technical Field
The invention belongs to the technical field of agricultural biological pesticides, and relates to application of chondroitin sulfate as an active ingredient in preparation of a plant immunity inducer.
Background
Plants are often affected by biological or non-biological factors during their growth and development, in morphology and in lifeChanges in physical and biochemical aspects, such as wilting, necrosis and deformity, seriously obstruct the normal growth of plants. Plant viruses are non-cellular organisms with simple structures and different forms, are parasitic in plants, and replicate and reproduce by using substances, energy and places provided by host cells. Because of limitations of the immune system of plants, control of plant viruses is difficult [1] . In agricultural production, the extent of viral disease is second only to fungi, a second major class of plant disease, also known as "plant cancer" [2] Is a key and difficult problem of plant disease research. Potato (Solanum tuberosum l.) is the largest non-cereal crop in the world, and is also the fourth largest crop worldwide next to wheat, rice and corn. The potato yield in China is high, the planting area is wide, and the potato is the biggest potato producer in the world. Potato late blight (Potato Late Blight) is a destructive oomycete disease caused by phytophthora infestans (Phytophthora infestans) and is listed as the first major disease of world food crops [3] . The disease causes about 20% yield reduction of potato in China, and can reach up to 50% in severe cases, even when the disease is dead [4] Causing serious economic loss and threatening food safety [5] . The chemical agent is an effective means for preventing and treating the diseases, can reduce the loss caused by the diseases, but is used for a long time and in a large range, and seriously pollutes the soil ecosystem and the surrounding ecological environment. Therefore, a novel disease-resistant strategy is searched, the disease-resistant mechanism of the plant is researched, the immunity of the plant is fundamentally improved, and the research and development of a novel environment-friendly disease-resistant agent for the disease has important application value and practical significance.
With the world population growing, there is an urgent need for creative and breakthrough strategies in controlling plant pathogens and increasing grain yield [6] . Plant immunity elicitors are a new class of pesticides that enhance plant disease and stress resistance by activating the immune system and regulating the metabolism of plants [7] . Plant immunity inducer itself has no direct bactericidal activity, and disease is prevented and treated mainly by promoting plants to utilize self natural immune system without depending on exogenous pesticideDirectly kills pathogens. The biogenic plant immunity inducer not only improves the disease resistance of plants, but also has the advantages of stimulating the growth of crops, improving the yield, improving the quality of agricultural products, having no residue and the like, being nontoxic to people and livestock, having simple processing technology, low cost and being not easy to generate drug resistance to germs, meeting the idea of realizing green prevention and control under the condition of effectively protecting the agricultural biodiversity, and having good development and application potential [8] 。
Chondroitin Sulfate (CS) is a complex, linear, anionic type of glycosaminoglycan covalently linked to proteins to form proteoglycans, widely distributed on the extracellular matrix and cell surface of animal tissues, the sugar chain consisting of alternating glucuronic acid and N-acetylgalactosamine (also known as N-acetylgalactosamine) disaccharide units linked to serine residues of core proteins through a sugar-like linking region [9] . Although the backbone structure of the polysaccharide is not complex, it exhibits a high degree of non-uniformity in terms of degree of sulfation, sulfate groups and distribution of the two different isomeric uronic acids within the chain. The functional region with a special structure in the polysaccharide chain specifically interacts with specific proteins to realize biological functions, functional regions with different structures exist in the same polysaccharide chain, and different proteins can interact to perform different functions. Functional regions of different structural sequences bind to specific proteins to perform specific functions. The fine structure of chondroitin sulfate determines the specificity of function and interaction with various protein molecules [10] . CS plays an important role in various normal physiological and pathological processes of animals, and shows various biological activities such as antioxidation, anti-atherosclerosis, antithrombotic, unobvious immunogenicity and the like. However, studies of CS in plant physiology and pathology have not been reported.
Chinese patent document CN108056101a (application No. 201610978760.9) discloses a chitosan-alginic acid nanoparticle, and preparation and application thereof, and the chitosan-alginic acid nanoparticle prepared by the patent document has activities of inducing plant resistance and promoting plant growth, but the application of chondroitin sulfate in plants is not involved in the patent document.
Disclosure of Invention
Aiming at the defects of the prior art, the invention provides application of chondroitin sulfate serving as an effective component in preparing a plant immunity inducer.
The invention aims to solve the problems of pesticide residue and environmental pollution caused by chemical pharmaceutical agents, and to discover a biological source antiviral agent which is nontoxic and residue-free and can protect people and livestock, and provides a novel application of chondroitin sulfate in inducing plant disease resistance.
The technical scheme of the invention is as follows:
the application of chondroitin sulfate as an active ingredient in the preparation of plant resistance inducer.
According to a preferred aspect of the present invention, the object to be elicited comprises a plant virus or a plant oomycete disease.
Further preferred plant viruses include tobacco mosaic virus, cucumber mosaic virus, potato virus Y, tomato yellow leaf curl virus, tomato stay green virus.
Further preferably, the plant oomycete disease comprises phytophthora infestans.
Preferably according to the invention, the plant comprises Nicotiana, cucumis, solanum, lycopersicon.
Further preferred, the plants include tobacco, cucumber, potato, tomato.
A plant resistance inducer contains chondroitin sulfate as effective component.
The application of the plant resistance inducer in plant cultivation.
According to a preferred aspect of the invention, the plant elicitor is used against plant viral and/or plant oomycete diseases.
According to a preferred aspect of the present invention, the chondroitin sulfate is used at a concentration of 20. Mu.g/mL to 55. Mu.g/mL.
It is further preferred that chondroitin sulfate is used at a concentration of 45-55. Mu.g/mL.
Advantageous effects
1. The invention provides a new application of Chondroitin Sulfate (CS) as a biological source immunity inducer in plant disease resistance, wherein the chondroitin sulfate can induce accumulation of plant endogenous salicylic acid; exciting active oxygen burst; improving the resistance of plants to pathogenic bacteria.
2. The Chondroitin Sulfate (CS) has good prevention effect at the use concentration of 25 mug/mL and 50 mug/mL, and the average prevention effects are respectively as follows: 76.02 percent and 85.84 percent of the composition are higher than the control 50 mug/mL Chitosan Oligosaccharide (COS) by 69.72 percent; the use cost of CS at 25-50 mug/mL is lower than that of Chitosan Oligosaccharide (COS) by comprehensive analysis, and the Chondroitin Sulfate (CS) provided by the invention is more economical to use.
Drawings
FIG. 1 is a graph showing the control effect and the quantitative view of potato X virus (PVX-GFP) with GFP fluorescence label inoculated after treatment with Chitosan Oligosaccharide (COS) as a positive control and with different concentrations of Chondroitin Sulfate (CS) as a negative control;
in the figure: a is a fluorescence intensity graph in the raw smoke after CS treatment with different concentrations;
b is qRT-PCR detection of GFP in the raw tobacco, and Bar value represents standard deviation, and P is less than or equal to 0.001.
FIG. 2 is a graph showing the effect of controlling phytophthora infestans inoculated after treatment with Chondroitin Sulfate (CS) and a quantitative graph of phytophthora infestans;
in the figure: a is a phenotype chart of the phytophthora infestans inoculated in the smoke after CS treatment;
b is the expression quantity of phytophthora infestans filaments in the smoke generated by combining a fungus detection kit with qRT-PCR, the data are derived from three biological experiments, and the Bar value represents standard deviation, and P is less than or equal to 0.001;
CK is sprayed ddH 2 The control group O, CS is a CS solution group sprayed with 50 mug/mL.
FIG. 3 is a graph showing the results of detection of Salicylic Acid (SA) content after Chondroitin Sulfate (CS) treatment and the expression levels of genes ICS and PAL;
in the figure: a is a content graph of SA in the smoke generated after CS spraying treatment;
b is an expression level diagram of SA synthetic genes ICS and PAL in the self-generated cigarettes after different treatments detected by qRT-PCR.
FIG. 4 is a graph showing DAB dyeing results of CS spraying treatment at different times;
in the figure: a is an accumulation diagram of CS promoting the hydrogen peroxide of the present smoke;
b is a graph of qRT-PCR assay gene CAT, SOD, APX, rbohA, rbohB results 24h after CS treatment, data expressed as mean (n=5) ±sd, < p <0.05, < p <0.01;
CK is water control group and CS is CS solution group of 50 μg/mL.
Detailed Description
The technical scheme of the invention is further described below with reference to the examples and the attached drawings, but the scope of the invention is not limited thereto.
It should be noted that the following detailed description is exemplary and is intended to provide further explanation of the invention. Unless defined otherwise, all technical and scientific terms used herein have the same meaning as commonly understood by one of ordinary skill in the art to which this invention belongs.
It is noted that the terminology used herein is for the purpose of describing particular embodiments only and is not intended to be limiting of exemplary embodiments according to the present invention. As used herein, the singular is also intended to include the plural unless the context clearly indicates otherwise, and furthermore, it is to be understood that the terms "comprises" and/or "comprising" when used in this specification are taken to specify the presence of stated features, steps, operations, devices, components, and/or combinations thereof.
Source of main materials
Chondroitin sulfate, chitosan oligosaccharide, was purchased from Qingdao Jiaweida biotechnology limited.
Example 1
Chondroitin Sulfate (CS) can improve virus resistance of plants
Firstly, uniformly stirring nutrient soil containing vermiculite and matrix (volume ratio is 1:2), and putting the nutrient soil into a disposable plastic small pot; 3-4 seeds of the tobacco are spread in each small pot, placed in a greenhouse (24 ℃,14h illumination and 10h darkness) and covered with a layer of plastic film for cultivation. After emergence, transplanting the seedlings into plastic small bowls, wherein each bowl is provided with 1 plant, and placing the seedlings in a greenhouse for management. The self-generated cigarettes at the age of 4 weeks are selected, and are sprayed with the medicament, 10 plants are sprayed for each treatment, and each plant is 5mL. 2 hours after the spraying of the medicament, potato virus X (PVX-GFP) inoculation liquid with GFP fluorescence labels is inoculated, friction inoculation is carried out, each plant is inoculated with 3 leaves, and each leaf is 50 mu L. The disease was investigated 5 days after inoculation.
The medicament treatments are 6 treatments of 25 mug/mL of chondroitin sulfate, 50 mug/mL of chondroitin sulfate, 100 mug/mL of chondroitin sulfate, 200 mug/mL of chondroitin sulfate, 50 mug/mL of chitosan oligosaccharide and clear water control, and each treatment is repeated for 3 times and is arranged randomly.
The preparation method of the virus comprises the following steps: collecting tobacco leaves infected with Potato Virus X (PVX), and treating fresh tobacco leaves according to the following conditions: buffer=1:20 g/mL weight/volume ratio and buffer, the type of buffer is PBS, and the pH value is pH7.0, so that PVX virus inoculation liquid is prepared.
Experimental results: phenotypes were observed (A in FIG. 1) and the content of PVX in the different treatments was detected using qRT-PCR (B in FIG. 1). The results showed that the average preventive effect at the CS use concentration of 25. Mu.g/mL and 50. Mu.g/mL were: 76.02% and 85.84% of the total chitosan oligosaccharide/COS compound is higher than that of the control 50 mug/mL by 69.72%.
Example 2
Chondroitin Sulfate (CS) enhances resistance of plants to Phytophthora infestans
In order to find out whether CS can enhance the resistance of plants to phytophthora infestans, the invention sprays 50 mug/mL CS solution to the self-generated cigarettes with consistent seedling age and same growth vigor and sprays ddH 2 O is a control, leaves at the same position are placed in a plastic dish after 24 hours, and the treated phytophthora infestans are inoculated to the same position of the leaves and repeated three times. After 3d, observing the disease conditions of different experimental groups, finding that the degree of withered and yellow of the CS treated smoke leaf is weaker, the area of the disease spots is smaller, and ddH 2 The degree of the yellow spot of the generated smoke treated by O is more serious, and the area of the disease spot is larger (A in figure 2), and the quantitative detection of the growth amount of hypha in the leaves of the generated smoke treated by different treatments is carried out by using a fungus quantitative kit and qRT-PCR. The result shows that the growth amount of phytophthora infestans filaments in CS pretreated tobacco leaves is obviously lower than ddH 2 The O-treated benthamic cigarette (B in FIG. 2), consistent with the phenotype, showed that CS was able to enhance plant resistance to Phytophthora infestans.
Example 3
Chondroitin Sulfate (CS) induces accumulation of salicylic acid
In the previous experiment, CS has been proved to promote plant disease resistance, 40 plants of benthames with consistent growth vigor and same seedling age are selected for exploring whether CS mediates plant disease resistance by promoting SA content, 20 plants are sprayed with CS solution of 50 mug/mL, and the other 20 plants are sprayed with ddH 2 O is used as a control, materials are obtained after 24 hours, the raw tobacco leaf slice extracts of sprayed water and CS are respectively extracted, and the content of Salicylic Acid (SA) is detected by a liquid chromatography, and the specific method is as follows:
(1) The extraction method comprises the following steps: 10.0g of leaf blades are fully ground and placed in a 50mL centrifuge tube, 4.0mL of 5% trichloroacetic acid is added, 30mL of diethyl ether is added after water is added to 20mL, the mixture is fully and evenly shaken, leaching is carried out for 12h, centrifugation is carried out for 5.0min under 1000g, the upper diethyl ether phase is taken out, the water phase is repeatedly extracted for 2 times by diethyl ether, diethyl ether phases are combined, vacuum rotary evaporation is carried out, 1.0mL of a mixed solution of 50% methanol and 50% acetic acid buffer (pH 3.2) is added for dissolution, and the mixture is placed in an Eppendorf tube for preservation, thus obtaining the free SA sample. Adding 18.5% HCl into the water phase to a final concentration of 3.2%, heating in 80 ℃ water bath for 1h, cooling, extracting with diethyl ether for 3 times, combining diethyl ether phases, evaporating to dryness, adding 1mL of a mixture of 50% methanol and 50% acetic acid buffer (pH 3.2), dissolving, and storing in an Eppendorf tube to obtain a combined SA sample. The bound SA sample was filtered through a 0.22 μm microporous filter and then detected by High Performance Liquid Chromatography (HPLC).
(2) Measuring content
The main instrument is as follows: and (3) an Shimadzu high performance liquid chromatograph.
SA standard, shanghai five-combined chemical plant production; methanol is chromatographic purity and is produced by Siyou biomedical technology Co., ltd; the remaining reagents used were all analytically pure.
HPLC detection conditions: c18 column, 7.3 mm. Times.20 cm; mobile phase methanol to acetic acid buffer (ph 3.2) =50:50; an Shimadzu fluorescence detector (excitation wavelength: 310nm, emission wavelength: 415 nm); the flow rate is 1mL/min; the sample loading was 20. Mu.L.
The results show that SA content in CS pretreated benthamic cigarettes is significantly higher than ddH 2 O pretreated control group shows that CS can promote the accumulation of SA in the cigarette and then promote pathogenResistance of bacteria (A in FIG. 3).
(II) studies have shown that plants synthesize SA through two pathways: the above experiments have demonstrated that CS is capable of promoting accumulation of SA in a raw smoke, and further define the signaling pathway upon which CS promotes an increase in the SA content of a raw smoke, by using the isochorismate synthase (Isochorismate Synthase, ICS) pathway and the Phenylalanine Ammonia Lyase (PAL) pathway, which use chorismate as a precursor.
Selecting 40 plants of self-generated cigarettes with consistent growth vigor and same seedling age, spraying 50 mug/mL CS solution on 20 plants, and spraying ddH on the other 20 plants 2 O is used as a control, materials are obtained after 24 hours, total RNA of the leaf of the present cigarette is extracted, and qRT-PCR is used for detecting related genes of the salicylic acid synthesis pathway.
The results showed that with ddH 2 The PAL gene expression level in CS-pretreated lamina of this tobacco leaf was significantly increased compared to the O-treated control group, whereas the ICS gene expression level was not significantly changed from the control group (B in fig. 3), indicating that CS pretreatment was able to synthesize salicylic acid via PAL pathway.
Example 4
Chondroitin Sulfate (CS) promotes accumulation of hydrogen peroxide
Reactive Oxygen Species (ROS) mainly include superoxide anions (O) 2 - ) Hydrogen peroxide (H) 2 O 2 ) And Nitric Oxide (NO), as a signaling molecule, is involved in the plant's defensive response to pathogens. When the Robh gene in the plant body is stimulated by the outside, the Robh gene rapidly causes the increase or decrease of the active oxygen through the activation or the inactivation of the Robh gene, and the Robh gene plays a role in the growth and the development of the plant and in biotic or abiotic stress.
In order to study whether CS affects the accumulation of ROS, the invention takes water treatment as a control, tobacco leaves of the same parts of different plants are taken down after 50 mug/mL CS is treated for 2H, 4H, 6H, 8H, 24H and 48H respectively, and the DAB staining method is used for evaluating H in the leaves 2 O 2 Is contained in the composition.
As can be seen from A in FIG. 4, a large amount of brown deposits were observed on the present smoke-producing lamina, indicating that CS induced H 2 O 2 DAB staining was preceded by accumulation of (C) over timeAnd weakening after strengthening. DAB was the deepest stain after CS 24H spraying, indicating that CS induced H 2 O 2 And reached a maximum after 24h of treatment.
Several ROS-scavenging enzyme-encoding genes at the RNA level, namely Catalase (CAT), superoxide dismutase (SOD), ascorbate (APX) and ROS-production-related genes, including respiratory burst oxidase homologous genes (RbohA and RbohB), were tested by qRT-PCR and tested before and after treatment. After 24h of CS treatment, the expression levels of CAT in plants were significantly down-regulated and the expression levels of RbohA and RbohB were significantly up-regulated compared to the control group (B in FIG. 4). This phenomenon suggests that accumulation of hydrogen peroxide is caused by the down-regulation of CAT expression and the up-regulation of RbohA and RbohB expression. These results indicate that CS can act as a plant immunity inducer to promote ROS outbreaks, thereby stimulating plant immunity to enhance its disease resistance.
The invention aims to solve the problems of pesticide residues and environmental pollution caused by chemical pharmaceutical agents, and discovers a biological source antiviral agent which is nontoxic and residue-free and can protect human and livestock, and provides a novel application of chondroitin sulfate as an immunity inducer in plant disease resistance. The results show that the CS has obvious effect when the concentration is 25-50 mug/mL, and the average prevention effect is respectively as follows: 76.02% and 85.84% of the total chitosan oligosaccharide/COS compound is higher than that of the control 50 mug/mL by 69.72%.
The inventor also performs an anti-plant virus experiment by using chondroitin sulfate from various sources, and the experimental effect is similar to that of the chondroitin sulfate, so that the chondroitin sulfate from different sources has the effect of inducing plant resistance.
Reference to the literature
[1] Chen Ji, shen Jiaxiang. Development of anti-plant virus agents and challenges and opportunities facing them. University of Yunnan agriculture journal 2005 (04): 505-512.
[2] Zhou Dehai, song Yuchuan, wang Yujun, zhu Changxiang. Test of efficacy of Cynanchum komarovii against tobacco viral diseases. Shandong agricultural science, 2007, (06): 76-78.
[3]Mladen Cucak RDAM(2021)Opportunities for Improved Potato Late Blight Management in the Republic of Ireland:Field Evaluation of the Modified Irish Rules Crop Disease Risk Prediction Model.Phytopathology 111:1349-1360.
[4] Zheng Gongmei the cause, symptoms and prevention of potato late blight modern animal husbandry science and technology 2019,11:44-45.
[5] Song Bafu, xie Kaiyun. Global late blight control initiative for CIP and participation in China. J.Potato, 1997,11 (1): 51-55.
[6]Gaffar FY,Koch A(2019)Catch Me If You Can！RNA silencing-based improvement of antiviral plant immunity.Viruses 11(7):673.
[7]Peng C,Zhang A,Wang Q,Song Y,Zhang M,Ding X,Li Y,Geng Q,Zhu C(2020)Ultrahigh-activity immune inducer from Endophytic Fungi induces tobacco resistance to virus by SA pathway and RNA silencing.BMC Plant Biology 20；169.
[8] Dewen the development and application of plant immunity inducer, chinese agricultural science and technology, 2014,16 (01): 39-45 [9]Sugahara K,Mikami T,Uyama T,Mizuguchi S,Nomura K,Kitagawa H (2003) Recent advances in the structural biology of chondroitin sulfate and dermatan sulfate. Current opinion in structural biology 13 (5): 612-620.
[10]Izumikawa,T.,Kitagawa,H.,Mizuguchi,S.,Nomura,K.H.,Nomura,K.,Tamura,J.,Gengyo-Ando,K.,Mitani,S.,and Sugahara,K.(2004).Nematode chondroitin polymerizing factor showing cell-/organ-specific expression is indispensable for chondroitin synthesis and embryonic cell division.J Biol Chem 279(51),53755-53761.
Claims (4)
1. The application of chondroitin sulfate as an active ingredient in preparing a plant resistance inducer, wherein the resistance inducer is an oomycete disease of plants, and the oomycete disease of plants is phytophthora infestans.
2. The use according to claim 1, wherein the plant is tobacco, cucumber, potato, tomato.
3. The use according to claim 1, wherein the chondroitin sulphate is used at a concentration of 20 μg/mL to 55 μg/mL.
4. Use according to claim 3, characterized in that the chondroitin sulphate is used in a concentration of 45-55 μg/mL.
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| JPS4828648B1 (en) * | 1970-09-11 | 1973-09-03 | ||
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Non-Patent Citations (1)
| Title |
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| ANTIVIRAL ACTIVITY OF POLYSACCHARIDES AGAINST INFECTION OF TOBACCO MOSAIC VIRUS;Yoh Sano;Macromol. Symp.;第99卷;239-242 * |
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