CN112400882A - Application of flagellin and/or chitohexaose in preparation of inhibitor for inhibiting photosynthesis of plants - Google Patents
Application of flagellin and/or chitohexaose in preparation of inhibitor for inhibiting photosynthesis of plants Download PDFInfo
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- CN112400882A CN112400882A CN202011424782.3A CN202011424782A CN112400882A CN 112400882 A CN112400882 A CN 112400882A CN 202011424782 A CN202011424782 A CN 202011424782A CN 112400882 A CN112400882 A CN 112400882A
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- 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
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- 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
- A01N63/00—Biocides, pest repellants or attractants, or plant growth regulators containing microorganisms, viruses, microbial fungi, animals or substances produced by, or obtained from, microorganisms, viruses, microbial fungi or animals, e.g. enzymes or fermentates
- A01N63/50—Isolated enzymes; Isolated proteins
Abstract
The invention provides application of flagellin and/or chitohexaose in preparing an inhibitor for inhibiting plant photosynthesis. The invention triggers the hypersensitive reaction of the plant by acting flagellin and/or chitohexaose on the pattern recognition receptor on the leaves of the plant, so that the cells around the infection point are rapidly programmed to die, thereby achieving the effect of inhibiting photosynthesis. The application of the invention is realized by causing ROS outbreak in the leaves of the plant, inducing PR gene expression, reducing PS II reaction center density and reducing NPQ of the leaves of the plant. The invention discloses application of a composition of flagellin and/or chitohexaose and a calcium compound in preparing an inhibitor for inhibiting plant photosynthesis.
Description
Technical Field
The invention belongs to the technical field of plant photosynthesis regulation, and particularly relates to application of flagellin and/or chitohexaose in preparation of an inhibitor for inhibiting plant photosynthesis.
Background
In the natural environment, plants are inevitably exposed to various microorganisms, some of which are able to reproduce themselves by infecting them. When infected, the plants accelerate the senescence and abscission of leaves, and the plants themselves prevent the further diffusion of pathogenic microorganisms, so that cells around the infected points are rapidly programmed to die, which is called as hypersensitive reaction and is a common mechanism of plant immune reaction. Plants recognize essential components conserved in microorganisms through a Pattern Recognition Receptor (PRR) to rapidly start an immune pathway, and the pathogenic microorganism conserved components recognized by the receptor are called disease course related molecular patterns (PAMPs), so that various stress-resistant signal molecules, such as calcium ions, active oxygen, various plant hormones and the like, are released.
The plant can utilize light energy to convert CO into chloroplast2And H2O is synthesized with organic matter for its vital activities and releases oxygen, a process known as photosynthesis. It is divided into light reaction and dark reaction, the light reaction mainly uses PS II and PSI composite protein to convert light energy into chemical energy, the dark reaction uses energy and substance produced by light reaction to convert CO2Conversion to organic material, this process is known as the calvin cycle. Plant photosynthesis is responsible for most of the oxygen production and biomass fixation on earth. Thus, when a plant is stressed, the plant cells are then stressed to a greater extent, in which case the physiological and biochemical parameters of the plant are necessarily reduced, which naturally also includes photosynthesis.
Disclosure of Invention
The invention aims to provide application of flagellin and/or chitohexaose in preparation of an inhibitor for inhibiting plant photosynthesis. The application of the invention can effectively cause plant immune response, thereby inhibiting plant photosynthesis and having good application prospect.
In order to achieve the purpose, the technical scheme of the invention is as follows:
the invention provides application of flagellin and/or chitohexaose in preparing an inhibitor for inhibiting plant photosynthesis.
Further, the flagellin and chitohexaose act on pattern recognition receptors on leaves of the plant to elicit an immune response.
Further, the flagellin and chitohexaose cause an ROS burst in the leaves of the plant.
Further, the flagellin and chitohexaose induce PR gene expression.
Further, the flagellin and chitohexaose decrease PS II reaction center density.
Further, Ca2+With the signal intact, immune regulation triggered by flg22 up-regulates the expression of the PS II peripherin genes PsbO and PsbP.
Further, the chitohexaose treatment caused a significant downregulation of both PsbO and PsbP.
Further, the chitohexaose-induced immune pathway leads to PsbS gene down-regulation as well as vde gene up-regulation.
Further, the flagellin and chitohexaose decrease the NPQ of the leaf of the plant.
Further, the plant is a thylakoid plant.
The invention also provides application of the flagellin and/or chitohexaose composition and a calcium compound in preparation of an inhibitor for inhibiting plant photosynthesis.
Further, the calcium compound is calcium nitrate tetrahydrate.
Compared with the prior art, the invention has the advantages and technical effects that: the flagellin and/or chitohexaose obviously promote the expression of plant leaf PR-4 and PR-10 genes, and simultaneously reduce the density of PS II reaction centers, reduce the PS I activity and reduce the energy dissipation of peanut leaves, thereby reducing the photoprotective ability of plants, achieving the effect of a photosynthesis inhibitor, having wide action range, high reliability and quick reaction in inhibiting photosynthesis.
The invention also provides Ca in calcium-containing compound for synergistically promoting and inhibiting photosynthesis2+Is the basic requirement of the plant growth process, and the calcium ions necessary for the plant growth process can have the promoting effect of inhibiting photosynthesis. The application provided by the invention is realized by triggering the hypersensitive reaction of the plant, so that the cells around the plant infection point are quickly programmed to die, and the senescence and the shedding of the peanut leaves are accelerated.
The herbicides used in modern agriculture mainly include four major classes, i.e., photosynthesis-inhibiting type, pigment synthesis-inhibiting type, amino acid biosynthesis-inhibiting type and lipid synthesis-inhibiting type. The flagellin and/or chitohexaose as photosynthesis inhibitor is very considerable in preparing biological source herbicide. And flagellin and/or chitohexaose have abundant resources, easy degradation, no residual toxicity, wide action objects, no damage to ecological environment, high economic benefit and good market application prospect.
Drawings
FIG. 1 shows the ROS content change in peanut leaves under flg22 treatment, A: peanut leaf H treated with 1. mu.M flg222O2The content of (c) is changed; b: peanut leaf O under 1. mu.M flg22 treatment2 -The content change of (c) note: *: the difference was significant compared to the control (p < 0.05).
FIG. 2 is the change in ROS content in peanut leaves under chitohexaose treatment, A: peanut leaf H treated with 200 mug/ml chitohexaose2O2The content of (c) is changed; b: peanut leaf O treated with 200 mug/ml chitohexaose2 -The content change of (c) note: *: the difference was significant compared to the control (p < 0.05).
FIG. 3 is a graph of the effect of flg22 treatment on pathogen-associated gene expression in peanut leaves, A: the expression condition of PR-4 genes of peanut leaves under flg22 treatment; b: the expression of PR-10 gene of peanut leaf treated by flg22 is shown as follows: *: the difference was significant compared to the control (p < 0.05).
FIG. 4 is the expression pattern of pathogen-associated genes in peanut leaves under chitohexaose treatment, A: the expression condition of PR-4 genes of peanut leaves under the treatment of chitohexaose; b: the expression condition of the PR-10 gene of the peanut leaf under the treatment of the chitohexaose is as follows: *: the difference was significant compared to the control (p < 0.05).
FIG. 5 is activity of reaction center per peanut unit under flg22 treatment, A: 1 μ M flg22 treatment of changes in ABS/RC of lower peanut leaves; b: 1 μ M flg22 treated peanut TROA change in/RC; c: 1 μ M flg22 treated peanut leaf ETOA change in/RC; d: DI peanut treated with 1. mu.M flg22OChanges in/RC, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 6 is activity of peanut unit reaction center under chitohexaose treatment, A: change in ABS/RC of peanut leaves treated with 200. mu.g/ml chitohexaose; b: TR of peanuts treated with 200. mu.g/ml chitohexaoseOA change in/RC; c: peanut leaf ET treated by 200 mu g/ml chitohexaoseOVariation of/RC D: DI peanut treated with 200. mu.g/ml chitohexaoseOChanges in/RC, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 7 shows the change in the reaction center density of peanut leaves under flg22 treatment, A: 1 μ M flg22 treated peanut leaf RC/CSO(ii) a change; b: 1 μ M flg22 treated peanut leaf RC/CSMChange, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 8 is the density change of peanut leaf reaction centers under chitohexaose treatment, A: peanut leaf RC/CS treated by 200 mu g/ml chitohexaoseO(ii) a change; b: peanut leaf RC/CS treated by 200 mu g/ml chitohexaoseMChange, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 9 shows Flg22 treated lower peanut leaf PI(abs)Change, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 10 is PI of peanut leaves under chitohexaose treatment(abs)Change, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 11 shows flg22 treated lower peanut leaves
The change of (2) note: *: the difference was significant compared to the control (p < 0.05).
FIG. 12 shows peanut leaves treated with chitohexaose
The change of (2) note: *: the difference was significant compared to the control (p < 0.05).
FIG. 13 is a flg22 treated lower peanut leaf PI(total)The change of (2) note: *: the difference was significant compared to the control (p < 0.05).
FIG. 14 is PI of peanut leaves under chitohexaose treatment(total)The change of (2) note: *: the difference was significant compared to the control (p < 0.05).
Fig. 15 is the energy dissipation change per unit area of peanut leaf under flg22 treatment, a: DI of peanut leaves treated with 1. mu.M flg22O/CSO(ii) a change; b: DI of peanut leaves treated with 1. mu.M flg22O/CSMChange, note: *: the difference was significant compared to the control (p < 0.05).
Fig. 16 is the heat dissipation variation per unit area of peanut leaf under chitohexaose treatment, a: DI of peanut leaves treated with 200. mu.g/ml chitohexaoseO/CSO(ii) a change; b: DI of peanut leaves treated with 200. mu.g/ml chitohexaoseO/CSMChange, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 17 is a graph of the change in NPQ of peanut leaves under flg22 treatment, notes: *: the difference was significant compared to the control (p < 0.05).
FIG. 18 is a graph of the change in NPQ of peanut leaves under chitohexaose treatment, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 19 is the relative expression levels of peanut PsbO and PsbP under flg22 treatment, A: the relative expression level of PsbO gene of lower peanut leaves treated by 1 mu M flg 22; b: the relative expression level of PsbP gene of lower peanut leaves treated by 1 mu M flg22 is shown as follows: *: the difference was significant compared to the control (p < 0.05).
FIG. 20 is the relative expression levels of peanut PsbO and PsbP under chitohexaose treatment, A: the relative expression level of PsbO gene of peanut leaves treated by 200 mu g/ml chitohexaose; b: relative expression level of PsbP gene of peanut leaves treated by 200. mu.g/ml chitohexaose is shown as follows: *: the difference was significant compared to the control (p < 0.05).
FIG. 21 is the relative expression levels of peanut PsbS and vde under flg22 treatment, A: the relative expression level of PsbS gene of lower peanut leaves is treated by 1 mu M flg 22; b: the relative expression level of vde gene of peanut leaves treated by 1 mu M flg22 is shown as follows: *: the difference was significant compared to the control (p < 0.05).
FIG. 22 is the relative expression levels of peanut PsbS and vde under chitohexaose treatment, A: the relative expression level of PsbS gene of peanut leaves treated by 200 mu g/ml chitohexaose; b: the relative expression level of the vde gene of the peanut leaves under the treatment of 200 mu g/ml chitohexaose is shown as follows: *: the difference was significant compared to the control (p < 0.05).
FIG. 23 is the relative expression levels of peanut CP12 under flg22 treatment, note: *: the difference was significant compared to the control (p < 0.05).
FIG. 24 is the relative expression levels of peanut CP12 under chitohexaose treatment, note: *: the difference was significant compared to the control (p < 0.05).
Detailed Description
The technical solution of the present invention is further described in detail with reference to the accompanying drawings and specific embodiments.
1. Laboratory apparatus
A continuous excitation type fluorometer (Handy PEA), a portable pulse modulation type fluorometer (FMS-2), an ultraviolet visible spectrophotometer (UV-1750), an ABI 7500FAST fluorescence quantitative PCR instrument, a refrigerated centrifuge, and the like.
2. Experimental reagent and kit
Flagellin flg22(sigma), chitohexaose (solarbio), potassium sulfate, magnesium sulfate heptahydrate, potassium dihydrogen phosphate, calcium nitrate tetrahydrate, disodium ethylene diamine tetraacetate, ferrous sulfate, boric acid, copper sulfate pentahydrate, zinc sulfate heptahydrate, ammonium bicarbonate, molybdic acid, acetone, etc. Total RNA extraction kit was purchased from Tiangen Biochemical technology Ltd, PrimeScriptTMRT reagent Kit with gDNA Eraser from TaKaRa, hydrogen peroxide (H)2O2) Reagent kit(H2O2-2-Y) and superoxide anion kit (SA-2-G) were purchased from Suzhou Keming Biotechnology, Inc.
Example 1: selection of PAMPs
The invention selects flagellin flg22(sigma) and chitohexaose (solarbio) as raw materials of the photosynthesis inhibitor, and researches the influence of an immune pathway initiated by the flagellin flg22(sigma) and the chitohexaose on the photosynthesis inhibition.
Flagellin flg22 (hereinafter referred to as flg22) belongs to the class of proteins PAMPs. Flagella are the motile organs of bacteria that enable bacteria to respond in the face of a stimulus. Flagellin is derived from the conserved N-or C-terminus of various bacterial flagella. Because of these characteristics, animal and plant cells recognize flagellin.
Chitosan is a chitosan, and belongs to saccharides PAMPs. Chitosan is a natural and biodegradable polymer material, has wide application in the fields of beauty, food, biology, medicine, agriculture and the like, and is deacetylated chitin. Like chitin, chitosan fragment, chitosan oligosaccharide, has biological effects as well as good water solubility, although it is a high polymer.
Example 2: leaf material culture
1. Experimental methods
(1) Selecting peanuts (Arachis Hypogaea, 25. rosette) with full seeds and similar sizes, soaking the peanuts in deionized water for 40 minutes, transferring the peanuts into a plastic white pot paved with wet gauze to ensure that the peanuts are not stacked, then adding another layer of wet gauze to provide a humid and breathable environment for the seeds, and placing the white pot into an incubator at 28 ℃ for two days to wait for germination.
(2) The germinated seeds are put into a small plastic basin (the upper caliber is 9cm, the lower caliber is 6.5cm and the height is 8cm) filled with quartz sand, the quartz sand is repeatedly washed by deionized water, the small plastic basin with a hole at the bottom is put into a white big basin to be covered with a film, and the big basin is transferred into a greenhouse for culture. The greenhouse is continuously illuminated for 14 hours every day in the daytime (400 mu mol m)-2s-1) The temperature was 25 ℃/20 ℃ (day/night). The membrane was removed by deionized water incubation for approximately 4 days.
(3) And dividing the peanut seedlings obtained in the last step into two groups, namely Calcium (CA) group and calcium-free (NC) group, pouring the Hoagland nutrient solution completely in the CA group, removing the calcium nitrate tetrahydrate in the Hoagland nutrient solution in the NC group, and balancing nitrogen by using ammonium bicarbonate. Before irrigation, the mother liquid of each part of nutrient solution is mixed and diluted to the final concentration, the pH value is adjusted to 6.5, and the large plastic basin is replaced with new nutrient solution every day.
(4) After 20 days of culture, taking peanut functional leaves, placing the leaves in vitro in a culture dish added with deionized water for dark adaptation for 4 hours, measuring relevant initial data of CA and NC groups, and respectively adding 1 mu M flg22 and 200 mu g/ml chitohexaose for treatment for 1 hour, 2 hours and 4 hours. And for measuring active oxygen and extracting RNA, the material is treated, sent to liquid nitrogen, frozen and stored at-70 ℃.
Example 3: fluorescence parameter determination
1. Experimental methods
(1) Chlorophyll rapid fluorescence kinetic parameter determination: after each group of isolated leaves was fully adapted to the dark using a continuous excitation fluorometer (Handy PEA), the leaves were clamped on a leaf clamp on a probe and measured in the dark throughout, and the fluorescence parameters are shown in Table 1.
TABLE 1 various fluorescence parameters
(2) Measurement of fluorescence quenching parameters: and (3) fully adapting each group of in-vitro blades in dark by using a pulse modulation type fluorescence instrument (FMS-2), clamping the blade clamp with the metal sheet, enabling the blade clamp with the blades to be close to the probe, and opening the metal sheet before measurement to operate a program.
Example 4: total RNA extraction
1. Experimental methods
Extracting total RNA of peanut leaves, and referring to the TIANGEN kit instruction.
(1) After placing the mortar in liquid nitrogen for precooling, fully grinding the peanut leaves by using the liquid nitrogen, taking 0.1g of ground powder, placing the ground powder into a 1.5ml centrifugal tube which is rotated by 1ml of lysis solution RZ, and placing the ground powder on an oscillator for a moment to ensure uniform mixing.
(2) The sample was allowed to stand at 25 ℃ for 5 min.
(3) The pre-set refrigerated centrifuge, which had reached 4 ℃, was adjusted to 12000rpm for 5min and after centrifugation the supernatant was aspirated as full as possible into another empty 1.5ml centrifuge tube.
(4) 200. mu.l of chloroform (unopened or used exclusively for RNA) was added thereto, and the mixture was shaken on a shaker for 15sec after capping and then allowed to stand at room temperature for 3 min.
(5) Centrifuging for 10min (same temperature and rotation speed as (3)), transferring the colorless aqueous phase into a new tube, and ensuring the purity and concentration of RNA in the later period by sucking the colorless aqueous phase into the middle layer and the yellow organic phase as much as possible.
(6) Adding 300 μ l anhydrous ethanol (pre-cooled in a refrigerator at 4 deg.C), mixing, transferring the obtained solution and precipitate into adsorption column CR3, centrifuging at 4 deg.C 12000rpm for 30sec, and discarding waste liquid.
(7) To the adsorption column CR3 was added 500. mu.l of deproteinized solution RD (ethanol had been added and the mixture was placed in a refrigerator at 4 ℃ for precooling), centrifuged for 30sec (at the same temperature and rotation speed as above), and the waste liquid was discarded.
(8) To the adsorption column CR3 was added 500. mu.l of the rinse-removing solution RW (to which ethanol had been added), followed by standing at room temperature for 2min, centrifugation for 30sec (temperature and rotation speed were the same as above), discarding the waste liquid, and repeating the operation once.
(9) Centrifuging for 2min (temperature and rotation speed unchanged), discarding the waste liquid, and air drying at a place close to the alcohol burner.
(10) The adsorption column CR3 was transferred to a new 1.5ml centrifuge tube, 50. mu.l of RNase-free ddH2O was added to the adsorption column film, and the mixture was allowed to stand at room temperature for 2min and then centrifuged for 2min (temperature and rotation speed were the same as above).
(11) The collected RNA was placed on ice for subsequent experiments or at-70 ℃ until use.
Example 5: RNA concentration and purity assays
Mu.l of the RNA obtained in example 3 were mixed with 49. mu.l of RNase-free ddH2O was mixed well in a new RNase Free centrifuge tube and the RNA concentration and purity were determined using an Eppendorf biophotometer.
Example 6: reverse transcription of RNA
1. Experimental methods
Reverse transcription of peanut RNA, PrimeScript referenced to TaKaRaTMRT reagent Kit with gDNA Eraser instructions.
(1) Removal of genomic DNA
Reagents were added to ice in the following order
Wherein Total RNA is the number of milliliters required for conversion based on the concentration of the extracted RNA obtained in example 3 and 2 mug as a standard;
after mixing, the mixture was reacted at 42 ℃ for two minutes and placed on ice.
(2) Reverse transcription reaction
Reagents were added to ice in the following order
Mixing, reacting at 37 deg.C for 15min, reacting at 85 deg.C for 5sec, and standing on ice or storing at-20 deg.C.
Example 7: real Time PCR reaction
1. Experimental methods
Table 2 shows the gene primer sequences, and the peanut TUA5 is used as an internal reference gene, and the specific steps are as follows:
(1) reaction system configuration
Prepared on ice in the following order
The cDNA concentration used for the RT PCR reaction was 5-fold diluted after reverse transcription in example 5.
(2) Setting of reaction program
Step 1: pre-denaturation at 95 ℃ for 30 sec;
step 2: PCR reaction, 40 cycles, 95 ℃, 5 sec; 60 ℃, 34 sec;
TABLE 2 Gene primer sequences
Example 8: active oxygen measurement
1. Experimental methods
With particular reference to hydrogen peroxide (H)2O2) Kits and superoxide anion (ORF) kit instructions.
(1) Effect of flg22 on the active oxygen content of peanut leaves
ROS play an important role in promoting tissue repair and combating pathogenic microorganisms in plants, often accompanied by a burst of reactive oxygen species in the early immune response of the plant. As can be seen from FIG. 1A, H in CA and NC peanut leaves after 1H and 2H treatment with flg222O2The rising amplitude of 1h is higher relative to other times, the active oxygen content begins to decrease, and the normal state is recovered at 4h, and the two trends are basically consistent. FIG. 1B shows O in flak 22-treated peanut leaves2 -The content was significantly increased and the content of CA treatment was significantly lower than NC treatment. The results show that: the immune response triggered by flg22 causes an outbreak of ROS, Ca, in peanut leaves2+To O2 -Has obvious inhibiting effect on the accumulation of the (D).
(2) Effect of chitohexaose on the active oxygen content of peanut leaves
After the treatment with chitohexaose, the active oxygen content of the peanut leaves was measured. As can be seen from FIG. 2A, peanut leaves treated with chitohexaose had H in the peanut leaves after 1H and 2H2O2All show a remarkable upward trend at 4h H2O2The content was reduced to about the initial level and there was no significant difference between the CA and NC treatments. As shown in FIG. 2B, the peanut leaves were treated with chitohexaose, O2 -The content was initially increased and then showed a downward trend, with no significant difference between CA and NC treatments. Shows that: the immune response triggered by chitohexaose causes a burst of reactive oxygen species in the peanut leaves.
2. Results of the experiment
flg22 and chitohexaose trigger the peanut immune pathway, and this process involves Ca2+And (4) signal path. Reactive Oxygen Species (ROS) are signal molecules in the immune process, Ca2+The active oxygen scavenging capacity of the peanut leaves is improved.
Example 9: plant leaf pathogen associated gene expression
1. Flagellin and chitohexaose triggered pathogen-associated gene expression
Plants, when exposed to PAMPs or pathogenic microorganisms, trigger an autoimmune response, in the downstream of which a mitogen-activated protein kinase (MAPK) cascade signal finally leads to a change in the expression of the pathogen-associated gene PR. The PR gene family is found in about 17 species in different plants, which can be induced by biotic or abiotic stress. In many cases, marker genes responsive to defense-related plant hormones such as salicylic acid, jasmonic acid and ethylene, the present invention selects for detection of PR-4 and PR-10 in the PR gene family. PR-4 is related to coding chitinase and has antifungal activity. The PR-10 gene family encodes small proteins with cytoplasmic localization that function as RNAses, post-translational modifications, etc., and respond to both abiotic and biotic factors.
(1) Expression pattern of pathogen-associated genes in peanut leaves treated with flg22
The PR gene may encode a pathogen-associated protein that is induced when a plant is infected with a pathogenic microorganism and participates in the defense response of the plant. As shown in figure 3, after the peanut seedlings in the CA group are treated by flg22, the PR-4 gene is significantly up-regulated at 1h, 2h and 4h, particularly the up-regulation of 1h and 2h is maximum, and 4h is moderate relative to 1h and 2 h. NC peanut seedlings are remarkably upregulated on 1 and 2hPR-4 genes after flg22 treatment, but the upregulation amplitude is obviously not as high as calcium, and is not remarkable after 4 hours. PR-10 gene expression of seedlings with or without calcium showed a significant increase, but the trend was relatively stable with calcium.
(2) Expression pattern of pathogen-associated genes in peanut leaves under treatment of chitohexaose
Treating peanut seedling leaves by using chitohexaose to detect pathogen-related genes. As can be seen from FIG. 4A, after peanut seedlings are subjected to chitosans, CA leaves are significantly upregulated in PR-4 and PR-10 genes at 1, 2 and 4h, peanut seedlings of NC are significantly upregulated in the chitosans-treated 4hPR-4 gene, PR-10 gene expression is significant at 4h, and NC is increased to a smaller extent relative to CA (FIG. 4B).
2. Results
flg22 and chitohexaose induced the expression of PR gene in peanut leaf and Ca2+The signal mediates the regulation and control of the expression of genes related to the peanut leaf pathogen by the chitohexaose, and the effect of stabilizing the immune response is achieved.
Example 10: activity of reaction center
1. Activity of PSII reaction center
(1) Activity per PS II reaction center after treatment with flg22
The antenna pigment in the green plant cell transfers the absorbed captured energy to the reaction center, and the reaction center uses the energy to reduce QAThus transferring electrons to the following photosynthetic reaction pathway, ABS/RC, TRO/RC、ETOReaction of three parameters,/RC, respectively representing the above processes, DIOthe/RC represents the energy dissipation per reaction center. From FIG. 5, it can be seen that ABS/RC, TRO/RC、DIOThe three parameters,/RC, have the same trend, CA treatment significantly rises at 1h and 4h after flg22 treatment, and NC significantly rises only at 4 h. ETOCA in the/RC only rises remarkably in 1h, 2h and 4h also have rising tendency but are not remarkable, and NC rises remarkably in 4 h. The results show that: the flg22 treatment increased the activity of PS II unit reaction centers of peanut leaves.
(2) Activity per PS II reaction center after chitohexaose treatment.
As can be seen in FIG. 6A, ABS/RC increased in both CA and NC after chitohexaose treatment; as can be seen from FIG. 6B, shell sixSugar treatment CA and NC TROboth/RC rises; FIG. 6C shows ET from chitohexaose-treated CA and NCOthe/RC rising tendency and ABS/RC and TROthe/RC is similar; FIG. 6D shows DI of chitohexaose treated NCOThe overall rise in/RC was insignificant and CA was significantly upregulated at 2 h. The results show that: after the chitosan treatment, the activity of PS II unit reaction center of peanut leaves is increased to a certain extent, but the activity is combined with Ca2+The signal relationship is not large.
2. Density of reaction center
(1) Reaction center density after flg22 treatment
Number of reaction centers in unit area of RC/CS reaction, RC/CSOAnd RC/CSMRespectively represent t ═ 0 and t ═ tFMCorresponding reaction center density. As can be seen from FIG. 7A, 1h after flg22 treatment, RC/CS was treated with CAOAnd a significant decrease, NC is 4h significant decrease. While in FIG. 7B, the CA processes the RC/CSMThe reduction is significant at 1h and 4h, and the reduction is significant at 4h in the NC group. The results show that: the immune response triggered by flg22 reduced the reaction center density, while a reduction in the number of reaction centers would certainly increase the burden on a single reaction center, which is responsible for the increased activity of a single PS II reaction center. In the pre-treatment period, the CA reaction center density was significantly lower than NC, since calcium responds positively to flg 22-triggered immune responses, mobilizing resources faster for the immune pathways, affecting reaction center density.
(2) Density of reaction center after treatment of chitohexaose
RC/CS of CA and NC after chitohexaose treatmentOSignificant downregulation occurred at both 2h and 4h (FIG. 8A), while RC/CSMThe CA group (as in fig. 8B) showed significant downregulation at 2 h. The results show that: chitosan-triggered immune responses reduce the reaction center density and increase the burden on individual reaction centers, but with Ca2+The signal paths are not very relevant.
3. Index of performance of light absorption energy of blade
(1) Performance index of light energy absorbed after flg22 treatment
PI(abs)Is an important parameter which can comprehensively reflect the density of the photo-responsive center, the light absorption and the electron transfer of the PS II.As can be seen from FIG. 9, the CA group of blades PI(abs)First decrease and then remain substantially stable, while the NC group remains substantially unchanged. This indicates that Ca is present during flg 22-induced immune pathway responses2+The signaling pathway reduces the reducing power of the PS II reaction center. Calcium ions are important ion signals of plant immune pathways, so that the CA immune pathway is triggered more rapidly than NC, and the influence range is wider.
(2) Index of performance of light energy absorbed after chitosan treatment
Unlike the previous flg22, the activity of the single reaction center is increased after treatment with chitohexaose, but the overall performance index PI of the leaf for absorbing light energy is increased(abs)And not changed (fig. 10). The results show that: chitosan-induced immune pathway was not directed against PI(abs)An influence is produced. The degree of immune response elicited by chitohexaose flg22 triggered immunity was evident by the relative expression of the PR genes, since chitohexaose was not like PI after flg22 treatment(abs)Creating a difference.
4. Results of the experiment
The immune response triggered by PAMPs can reduce the density of the reaction center and increase the activity of PS II unit reaction center of peanut leaves. In the early stage of treatment, since calcium responds positively to flg 22-triggered immune responses, and resources can be mobilized more quickly for immune pathways, the CA reaction center density is significantly lower than NC, and CA is present in2+The response speed of the part can be accelerated.
Ca during PAMPs-induced immune pathway response2+The signaling pathway reduces the reducing power of the PS II reaction center. Calcium ions are important ion signals of plant immune pathways, so that the CA immune pathway is triggered more rapidly than NC, and the influence range is wider.
Example 11 relative Activity of PS I
1. Quantum yield of PSI acceptor side terminal electron acceptor
As can be seen from FIG. 11, 1. mu.M flg22 treatment of 1h and 2h, CA leaves
Significantly reduced, and NC at 2h
The significant reduction, the remaining treatment time was not significant. The results show that: immune regulation induced by flg22 decreased the relative activity of PS I, while Ca2+The response speed of (2) seems to be more positive.
FIG. 12 shows
CA and NC were not significantly changed before and after chitohexaose treatment. The results show that: chitohexaose treatment did not affect the receptor side quantum yield of PS I from peanut leaves.
2. Index of comprehensive properties
PI(total)Is a parameter for integrating the electron transfer ability of the reaction from the energy of PS II to the acceptor side terminal of PS I. As can be seen from FIG. 13, PI of CA leaf was treated at 1. mu.Mflg 22 for 1h, 2h and 4h(total)Significantly reduced, and NC at 2h PI(total)The significant reduction, the remaining treatment time was not significant. The results show that: the immune regulation induced by flg22 reduced the photosynthetic activity of peanut leaves, while Ca2+The response is more positive.
As can be seen from FIG. 14, PI of CA and NC groups before and after chitosans treatment of peanut leaves(total)Not significant. The results show that: chitosan treatment does not affect the comprehensive performance index PI(total)。
Example 12 energy dissipation
During photosynthesis, the captured light energy is consumed primarily by photochemical electron transfer, chlorophyll fluorescence emission, and heat dissipation. Photochemical electron transfer is involved in the synthesis of photosynthetic products, and chlorophyll fluorescence emission is only a small part of the light energy consumption. Excess energy can damage PS II, and therefore, heat dissipation is an important way to consume excess light energy and prevent photodamage. The excess excitation energy is dissipated in a harmless manner as a process called non-photochemical quenching (NPQ). Typically, high intensity illumination or reduced efficiency of reaction centers increases energy dissipation.
Recombination of the PS II reaction center complex is a common mechanism of NPQ, including reversible inactivation and synthetic regeneration of the D1 protein, Ca2+The content of D1 protein in the treated plant is high. Turnover of protein components in the center of photosynthetic reactions is regulated by calmodulin (CaM), Ca2+Important components of signal transduction pathways. The Arabidopsis thaliana mutant VDE activity deletion mutant npq1, in npq1, prevented zeaxanthin accumulation due to point mutations in the VDE. Whereas the Arabidopsis thaliana mutant PsbS deletion mutant NPQ4, zeaxanthin can accumulate, but NPQ4 is also quench-inactivated, NPQ requires the simultaneous presence of VDE and PsbS due to interaction between VDE and PsbS, and PsbS acts at the quenching initiation site. The synthesis of Violaxanthin Decyloxidase (VDE) is subject to Ca2+Influence of (1), Ca2+Under the condition, CaM can mediate the expression of VDE genes and promote the circulation of lutein.
1. Heat dissipation per unit area
DIO/CSOAnd DIO/CSMRespectively represent t ═ 0 and t ═ tFMCorresponding heat dissipation per unit area. It can be seen from FIG. 15 that DI was only performed by NC at flg22 for 4hO/CSOAnd DIO/CSMThe decrease was significant, while neither CA was significant. The results show that: ca2+The heat dissipation per unit area of peanut leaves under the flg22 treatment was not greatly affected.
As can be seen in FIG. 16, in the group CA, DI after treatment with chitohexaoseO/CSOSignificant downregulation at 2h and 4h, DIO/CSMSignificant downregulation occurred at 2 h; in NC group, DIO/CSOSignificant downregulation occurred at 4 h. The results show that: chitosan treatment results in a reduction in heat dissipation per unit area, but with Ca2+The effect of (2) is not great.
2. Non-photochemical quenching
NPQ consumes excess energy from plants, and is associated with the PsbS protein and lutein cycle, consisting ofAs can be seen in FIG. 17, the NPQ of the CA leaf was significantly reduced for the 1 μ M f g22 treatments 2h and 4h, while the NPQ of the NC was higher than the CA leaf. The results show that: ca2+Involved in reducing non-photochemical energy dissipation during flg 22-stimulated immune responses.
The above results show that: the immune pathway induced by flg22 reduced the energy dissipation of peanut leaves, and the down-regulation of NPQ required Ca2+And (4) participating.
As can be seen in FIG. 18, the NPQ of chitohexaose-treated CA leaf was significantly lower than that of NC leaf. The results show that: chitosan-induced immune response Process, Ca2+Participate in the down-regulation of the NPQ of the peanut leaves.
The above results show that: the chitohexaose-induced immune pathway reduces the energy dissipation of the peanut leaves. This is essentially consistent with the results of flg22 treatment, indicating that non-photochemical energy dissipation is not the only mechanism for heat dissipation during the course of an immune response.
Example 13: effect of photosynthesis-related Gene expression
1. Relative expression level of PsbO and PsbP genes of peanut leaves
In higher plants, PS II is a multi-subunit complex mounted on the thylakoid membrane and possesses its own specific light-harvesting system consisting of two chlorophyll a binding proteins CP43(PsbC) and CP47(PsbB) and several chlorophyll a/b binding proteins, the most abundant of which is light-trapping complex II (LHC-II). PSII uses solar energy to catalyze a series of electron transduction reactions and causes the splitting of water molecules into oxygen, protons and electrons. The hydrolysis reaction is mainly catalyzed by the oxygen evolving complex, which is protected by the peripherins attached to the luminal side of PS II, which are PsbO, PsbP and PsbQ, respectively. PsbO is almost present in the photosynthetic organisms used, whereas PsbP and PsbQ are only found in green algae and plants, while in blue algae they are replaced by PsbV and PsbU, respectively. Participation of PsbO in maintaining Mn4CaO5Stability of the clusters, Studies show Mn4CaO5Clusters are a necessary prerequisite for water oxidation, the oxygen evolving compound Mn4CaO5Being chemically unstable as a free cofactor, it must be assembled on the photosystem II oxygen evolving complex at the energy gained by charge separation. Is bornIn the process of substance generation, natural inorganic cofactor Mn2+、Ca2 +And Cl-Binds to the newly translated PSII protein and assembles into an oxygen evolving complex through a series of light and shade steps. While PsbQ and PsbP require Cl-And Ca2+As an essential cofactor, Ca2+And also participates in the s-state cycle related to water decomposition. Hydrolysis to generate electrons which are transported to Nicotinamide Adenine Dinucleotide Phosphate (NADP)+) NADPH is generated by linear electron flow for the calvin cycle.
(1) Relative expression of PsbO and PsbP genes after flg22 treatment
PsbO and PsbP are PS II peripherins, and are important in the functional stability of PS II. From FIG. 19A, the PsbO gene of CA and NC leaves was significantly up-regulated at 1h and 4h, while the NC was more elevated. In FIG. 19B, the PsbP gene is specifically the same trend. The results show that: ca2+With intact signal, flg 22-triggered immunomodulation upregulated the expression of the PS II peripherin genes PsbO and PsbP, while Ca2+In the absence of signal, the up-regulation amplitude is larger, which promotes PS II repair faster.
(2) Relative expression of PsbO and PsbP genes after chitohexaose treatment
As can be seen from fig. 20, CA group PsbO decreased significantly at 1h after chitohexaose treatment, whereas NC decreased significantly at 4 h; in fig. 20B, CA groups were significantly reduced by chitohexaose treatment for 1h and 4h PsbP, while NC groups were significantly reduced for 1h, 2h and 4 h. The results show that: chitohexaose treatment caused significant downregulation of both PsbO and PsbP, with and without calcium. From the previous parameters, it can be seen that chitohexaose-triggered immunomodulation is less pronounced than flg22, and that no significant difference in partial photosynthetic parameters occurs, indicating that chitohexaose treatment does not have much influence on the PS II peripherin PsbO and PsbP, and therefore does not require accelerated PS II protein repair.
2. Relative expression level of PsbS and vde genes of peanut leaves
(1) Relative expression of PsbS and vde genes after treatment
The PS II protein PsbS and Violaxanthin Decyloxidase (VDE) are non-photochemically quenching (NPQ) components. FIG. 21A shows that NC treated 1hPsbS significantly at flg22Up-regulated, whereas CA leaves were significantly down-regulated at 2 h. Fig. 21B, CA was significantly downregulated at 2h, NC leaves were significantly downregulated at 1h and 2 h. Since they are all related to NPQ, both CA groups were down-regulated at 2h, while both NC groups were essentially complementary, suggesting that Ca is a major factor2+In the intact case, NPQ was inhibited in the flg 22-triggered immune response.
(2) Relative expression of PsbS and vde genes after chitohexaose treatment
Figure 22A shows that both the CA and NC groups PsbS genes were significantly downregulated after chitohexaose treatment and the difference was not significant. Fig. 22B shows significant upregulation of NC leaves 1h and 4h, while a significant and more significant increase in CA vde gene expression occurred. The results show that: the chitohexaose-induced immune pathway leads to PsbS gene down-regulation and vde gene up-regulation, while Ca2+The expression regulation and control effect on the vde gene is more obvious.
3. Peanut leaf CP12 gene relative expression level
(1) Relative expression of CP12 Gene after treatment with flg22
CP12 is a nucleic acid-encoded chloroplast protein that interacts with glyceraldehyde-3-phosphate dehydrogenase to participate in the Calvin cycle. As can be seen from fig. 23, CA was significantly down-regulated at flg22 treatment for 1h and 2h for CP12, but was significantly up-regulated at 2h and 4h for NC treatment. The results show that: in the course of an immune response, Ca2+When present, down-regulates the expression of CP12 gene, while Ca2+In the absence, expression of the CP12 gene is induced.
(2) Relative expression of CP12 Gene after Chitosan treatment
As can be seen in fig. 24, significant downregulation occurred in CA group 4h CP12 after chitohexaose treatment, and significant downregulation of 2h and 4h of NC leaf discs. The results show that chitohexaose-induced immunomodulation inhibits CP12 gene expression, while Ca2+Slowing down this effect.
Through the above experiments, the effect of flagellin and chitohexaose on inhibition of photosynthesis in plants can be obtained. As a photosynthesis inhibitor, flagellin and chitohexaose can be effectively combined with recognition receptors on plant leaves to induce plants to generate immune response, and influence on expression and energy dissipation of related genes for photosynthesis of the plants, so that the purpose of inhibiting photosynthesis is achieved, and the preparation method has good market application prospect in preparation of biological source herbicides.
The above examples are to be construed as merely illustrative and not limitative of the remainder of the disclosure. Although the present invention has been described in detail with reference to the foregoing embodiments, it will be apparent to those skilled in the art that various changes may be made and equivalents may be substituted for elements thereof; and such modifications or substitutions do not depart from the spirit and scope of the corresponding technical solutions.
Claims (9)
1. Use of flagellin and/or chitohexaose for the preparation of an inhibitor for inhibiting photosynthesis in a plant.
2. Use of flagellin and/or chitohexaose according to claim 1 for the preparation of an inhibitor for inhibiting photosynthesis in plants, wherein: the flagellin and chitohexaose act on pattern recognition receptors on the leaves of the plant, causing an immune response.
3. Use of flagellin and/or chitohexaose according to claim 1 for the preparation of an inhibitor for inhibiting photosynthesis in plants, wherein: the flagellin and chitohexaose cause an ROS burst in the leaves of the plant.
4. Use of flagellin and/or chitohexaose according to claim 1 for the preparation of an inhibitor for inhibiting photosynthesis in plants, wherein: the flagellin and chitohexaose induce PR gene expression.
5. Use of flagellin and/or chitohexaose according to claim 1 for the preparation of an inhibitor for inhibiting photosynthesis in plants, wherein: the flagellin and chitohexaose reduce the density of PSII reaction centers.
6. Use of flagellin and/or chitohexaose according to claim 1 for the preparation of an inhibitor for inhibiting photosynthesis in plants, wherein: the flagellin and chitohexaose reduce the NPQ of the leaf of the plant.
7. Use of flagellin and/or chitohexaose according to any of claims 1-6 for the preparation of an inhibitor for inhibiting photosynthesis in plants, wherein: the plant is a thylakoid plant.
8. Use of a combination of flagellin and/or chitohexaose as claimed in claim 1 and a calcium compound for the preparation of an inhibitor for inhibiting photosynthesis in plants.
9. Use of a combination of flagellin and/or chitohexaose with a calcium compound according to claim 8 for the preparation of an inhibitor for inhibiting photosynthesis in plants, characterized in that: the calcium compound is calcium nitrate tetrahydrate.
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| CN102242171A (en) * | 2011-05-04 | 2011-11-16 | 北京农学院 | Flagellin extract and novel purpose thereof |
| CN105076135A (en) * | 2015-09-22 | 2015-11-25 | 中国科学院海洋研究所 | Chitosan oligosaccharide regulator for promoting photosynthesis and growth of wheat |
| CN111491654A (en) * | 2017-07-20 | 2020-08-04 | 斯波根生物技术公司 | Biologically active polypeptides for improving the protection, growth and productivity of plants |
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| CN102242171A (en) * | 2011-05-04 | 2011-11-16 | 北京农学院 | Flagellin extract and novel purpose thereof |
| CN105076135A (en) * | 2015-09-22 | 2015-11-25 | 中国科学院海洋研究所 | Chitosan oligosaccharide regulator for promoting photosynthesis and growth of wheat |
| CN111491654A (en) * | 2017-07-20 | 2020-08-04 | 斯波根生物技术公司 | Biologically active polypeptides for improving the protection, growth and productivity of plants |
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| 郭卫华 等: "壳寡糖对烟草幼 苗生 长和光合作用及与其相关生理指标的影响", 《植物生理学通讯 》 * |
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