CN114158417B - Application of 4-methyl umbelliferone in improving drought resistance of plants - Google Patents
Application of 4-methyl umbelliferone in improving drought resistance of plants Download PDFInfo
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01G—HORTICULTURE; CULTIVATION OF VEGETABLES, FLOWERS, RICE, FRUIT, VINES, HOPS OR SEAWEED; FORESTRY; WATERING
- A01G17/00—Cultivation of hops, vines, fruit trees, or like trees
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01G—HORTICULTURE; CULTIVATION OF VEGETABLES, FLOWERS, RICE, FRUIT, VINES, HOPS OR SEAWEED; FORESTRY; WATERING
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Abstract
The invention discloses application of 4-methyl umbelliferone in improving drought resistance of plants, and finds that 4-MU can promote root system development of plants, particularly Malus plants, improve photosynthesis of plant leaves under drought stress, improve water utilization efficiency and further promote drought resistance of the Malus plants. Can promote the sustainable development of the apple industry in arid areas to a great extent.
Description
Technical Field
The invention belongs to the field of plant planting, and particularly relates to application of 4-methyl umbelliferone in improving drought resistance of plants.
Background
In recent decades, the fruit tree industry in China has achieved huge achievements, so that the economic benefits are increasing day by day, and the fruit tree industry becomes an important support for rural economic development in partial areas. Apple is one of the fruit trees with the longest cultivation history in the world, and wins the favor of people of all countries by the unique flavor and taste. The unique geographical location and climatic conditions of the high loess area become one of the largest and optimal apple producing areas in the world, but the areas have less rainfall and uneven seasonal distribution, so that the areas suffer continuous drought throughout the year. Drought is one of the most important limiting factors affecting the growth and development of fruit trees. Drought occurs in early summer, can affect the flowering and fruit setting of fruit trees, and has great influence on the yield of the fruit trees; drought occurs after fruit picking, which affects the flowering and fruit setting of fruit trees in the next year and further affects the fruit tree yield; drought stress can destroy the tissue structure of plants, reduce the water content and chlorophyll content of plant leaves, destroy the transmission of photosynthetic electrons, and influence the photosynthesis of plants; at the same time, drought stress also affects fruit quality characteristics such as size, color, sugar, acidity, hardness, etc.
In recent years, with the rapid advance of modern biotechnology, the production and living styles of people are continuously influenced. Metabonomics, an emerging biotechnology, has been widely used in various fields of plant research. The application of metabonomics in the field of fruit trees can comprehensively understand metabolites of fruit trees in various environments, perform quantitative analysis, and perform research on various aspects such as quality evaluation, fine variety breeding, plant stress tolerance response and the like. The plant metabolites not only provide human with necessary living products such as nutrition, energy, medicines and the like, but also are indispensable in the growth and development process of plants, and simultaneously, the plant metabolites also play important roles in the aspect of resisting biotic and abiotic stresses of the plants.
The root system is the most direct organ for sensing the soil moisture state and transmits signals upwards to various tissues and organs. Under drought stress, the root system can generate a series of morphological and structural changes, and the drought tolerance of the fruit tree is improved by increasing the root cap ratio to absorb more water and nutrients. Therefore, root metabolites are researched through metabonomics, metabolites capable of improving drought resistance of apple trees are excavated, functions and action mechanisms of the metabolites are researched, and sustainable development of apple industry in loess plateau regions can be promoted to a great extent.
Disclosure of Invention
The technical problem to be solved by the invention is as follows: how to excavate metabolites capable of improving drought resistance of apple trees.
The technical scheme of the invention is as follows: use of 4-methylumbelliferone for improving drought resistance in plants.
Further, the plant is an Malus plant.
Furthermore, the improvement of the drought resistance of the plant refers to the promotion of root system development, the improvement of leaf photosynthesis or/and the improvement of water utilization efficiency.
Analysis of metabolome data shows that after natural drought treatment is carried out on the drought-resistant stock, namely, the wild apple in Xinjiang, the contents of 4-methylumbelliferone (4-MU) and 4-methylumbelliferone-beta-D-glucoside (4-MU-Glc) are obviously increased; while the metabolite content change could not be detected in the drought sensitive rootstock T337 root system. Therefore, 4-MU as a precursor of 4-MU-Glc may play an important role in drought resistance of Malus plants. Selecting small Xinjiang wild apples and T337 seedlings, transplanting the seedlings into a flowerpot, applying 4-MU (0, 125 and 500 MU M) with different concentrations to roots of the small Xinjiang wild apples, and carrying out long-term natural drought treatment (60 days) after the small Xinjiang wild apples and the T337 seedlings naturally grow for 40 days.
Compared with the prior art, the invention has the following effects:
the invention discovers that 4-MU can promote the root development of plants, particularly Malus plants, improve the photosynthesis of plant leaves under drought stress, improve the water utilization efficiency and further promote the drought resistance of the Malus plants. Can promote the sustainable development of apple industry in arid areas to a great extent.
Drawings
FIG. 1: analysis of drought-differential metabolites in response to Sinkiang Malus domestica and T337. A: and (5) short-term natural drought treatment. B: differential metabolite detection. D _ T337: t337 naturally drought processing of the root system sample; c _ T337: t337 normal water supply root system sample; d _ XJ: carrying out natural drought treatment on a root system sample of the Malus sieversii; c _ XJ: samples of the normal water supply root system of the Xinjiang wild apple.
FIG. 2:4-MU influences the growth and development of Malus sieversii under long-term drought stress. A: phenotypic characteristics of Sinkiang Malus sieversii under different treatment conditions. B: the plant height is high. C: the stem is thick. D: dry weight of aerial parts. E: root dry weight. F: root-crown ratio.
FIG. 3: effect of 4-MU on growth and development of T337 under long-term drought stress. A: t337 phenotypic characteristics under different treatment conditions. B: the plant height is high. C: the stem is thick. D: dry weight of aerial parts. E: root dry weight. F: root-crown ratio.
FIG. 4: leaf relative water content. A relative water content of leaves of Malus sieversii. Bt 337 relative moisture content of leaf.
FIG. 5:4-MU influence on photosynthesis index of Sinkiang Malus domestica under long-term drought stress.
FIG. 6: influence of 4-MU on T337 photosynthesis index under long-term drought stress.
FIG. 7: influence of 4-MU on water utilization efficiency of Malus sieversii and T337 under long-term drought stress. A: instantaneous water utilization efficiency of Xinjiang wild apples. B: t337 instantaneous water use efficiency. C: carbon isotope (delta) of Malus sieversii13C) The abundance ratio. D: t337 Stable carbon isotope (. Delta.)13C) The abundance ratio.
Detailed Description
The experimental procedures in the following examples are all conventional ones unless otherwise specified. The test materials used in the following examples were all commercially available unless otherwise specified.
An apple root metabolite, 4-methylumbelliferone, can promote the drought resistance of apple plants under long-term drought stress by promoting root development and improving photosynthesis of plants. The test method and conclusion are as follows:
1. materials and methods
1.1 test materials and methods
The plant materials are drought-resistant stock, xinjiang wild apple and drought-sensitive stock T337, and are transplanted to a five-spring comprehensive test station of northwest agriculture and forestry science and technology university.
Short-term natural drought treatment: selecting drought-resistant anvil Xinjiang wild apples (XJ) and drought-sensitive anvil T337 (T337) of apples as test materials, transplanting the test materials into a field for pot culture test, and randomly dividing the test materials into 2 groups (5 pieces/group): control group (normal water supply group) and test group (natural drought treatment group). And (3) irrigating the potted plant to a saturated state, normally supplying water in the growth process of the control group, and stopping supplying water to the test group. After 20 days of natural drought treatment, the roots of the plant samples were sampled and stored at-80 ℃ for future use.
Metabolome assays: taking a root sample subjected to short-term natural drought treatment, grinding by using liquid nitrogen, weighing 100mg of powder, transferring the powder into a 1.5ml centrifuge tube, adding 120 mu l of 50% methanol, fully and uniformly mixing by vortex oscillation, standing for 10min, transferring to a refrigerator at minus 20 ℃ for overnight treatment to precipitate protein in the sample, transferring to a low-temperature centrifuge for 4000g for centrifugation for 20min, transferring supernatant to a 96-well plate, and performing non-target metabolite detection by using a high-resolution mass spectrometer (TripleTOF 5600plus, SCIEX).
4-MU treatment: selecting plant materials with more consistent growth vigor, transplanting the plant materials to a field pot for growing, continuing to grow for 2 weeks after new leaves grow out, selecting the plant materials with more consistent growth vigor again, applying 4-MU (0, 125 and 500 MU M) with different concentrations, and randomly dividing the plant materials into 2 groups after the plant materials grow for 30 days: control (Control) and long-term Drought-treated (Drought), 15 per group.
And (3) long-term drought treatment: and (3) combining a TDR soil moisture meter with a weighing method, and performing quantitative water control treatment. Control group: the volume water content of the soil measured by TDR is 43-48%, and the water content of the soil measured by a weighing method is 70-100%; long-term drought treatment group: the volume water content of the soil measured by TDR is 18-23%, and the water content of the soil measured by weighing method is 35-50%. The calculation method for measuring the water content of the soil by a weighing method comprises the following steps: soil water content = (soil weight-dry soil weight)/(soil water-saturated weight-dry soil weight).
Photosynthetic index determination: photosynthetic indicators including photosynthetic rate (Pn), stomatal conductance (Gs), intercellular CO2 concentration (Ci), and transpiration rate (Tr) were measured on plants using photosynthetic apparatus LI-COR 6400. When the photosynthetic index is measured, the leaves with relatively consistent tender degree, size and the like (the 7 th to 9 th leaves from top to bottom) are selected at 9-11 am with good weather for measurement.
Measuring the water content of the leaves: picking the leaves with relatively consistent tender degree and size (5 th to 7 th leaves from top to bottom), and recording the fresh weight M1 of the leaves; soaking the leaves in purified water for 24 hours, taking out the leaves, sucking water by absorbent paper, and recording the water saturation weight M2 of the leaves; and finally, transferring the leaves to an oven at 80 ℃ to dry the leaves to constant weight, and recording the drying weight M3 of the leaves. Leaf relative water content (M) = (M1-M3)/(M2-M3).
Measuring the water utilization efficiency: including photosynthesizing method and stable carbon isotope method (delta)13C) And so on.
The photosynthetic apparatus method can measure Instantaneous Water Use Efficiency (iWUE) = photosynthetic rate (Pn)/transpiration rate (Tr).
Water Use Efficiency (WUE) by stable carbon isotope assay: after the long-term drought continuous treatment for 60 days, collecting mature leaves (7 th to 9 th leaves from top to bottom) with relatively consistent tender degree and size on the ground, and wiping dust on the surfaces of the leaves by using a paper towel; randomly repeating 3 biological processes for each treatment, transferring to a 105 deg.C oven, deactivating enzyme for 30min, adjusting temperature to 80 deg.C, and oven drying to constant weight; transferring the dried sample to a ball mill, grinding, sieving with 80 mesh nylon gauze, collecting sample powder, and measuring with stable carbon isotope mass spectrometer (Flash EA 1112HT-Delta V Advantages, thermo Fisher)13C/12The abundance ratio of C.
2. Results and analysis
2.1 analysis of differences in metabolites of Sinkiang Malus sieversii and T337 root systems in response to external drought
Drought stress can induce plants to generate a series of metabolites to achieve a defense effect, and the root system is the most direct organ for sensing the soil moisture state, so that the metabolite difference of the root system under the drought stress is revealed, and a new insight is provided for screening the drought-resistant metabolites.
In the test, drought-resistant apple rootstock Xinjiang wild apple and drought-sensitive rootstock T337 are used as test materials, normal water supply and natural drought treatment are carried out (A in figure 1), root samples are taken for differential metabolite detection, and the results show that: detecting 131 different metabolites in a drought treatment group and a control group of root systems of Malus sieversii; 180 differential metabolites were detected in the T337 root drought-treated group and the control group (B in fig. 1). Metabolome data analysis showed that: 106 kinds of drought-shared differential metabolites are responded by root systems of Malus sieversii and T337; the root system of Malus sieversii responds to 25 drought-specific metabolites, of which the contents of 4-methylumbelliferone (4-MU) and 4-methylumbelliferone-beta-D-glucoside (4-MU-Glc) are significantly increased, while no change in the content of the metabolites can be detected in the drought-sensitive rootstock T337 root system. Therefore, 4-MU as a precursor of 4-MU-Glc may play an important role in the drought-resistant process of Malus plants.
2.2 differential metabolite 4-MU functional validation
2.2.1 4-MU phenotypic analysis of plant growth under Long-term drought stress
Drought stress can induce the generation of metabolites of 4-MU and 4-MU-Glc of the root system of Xinjiang wild apple which is a drought-resistant stock, and the existence of the metabolites cannot be detected in the root system of T337 of drought-sensitive stock. 4-MU can generate 4-MU-Glc under the catalytic action of UDP-glucosyltransferase (UGTs), and whether the 4-MU plays an important role as a precursor of the 4-MU-Glc in the drought resisting process of the Xinjiang wild apple is determined, the Xinjiang wild apple and T337 are adopted for long-term drought treatment, 4-MU (0, 125 and 500 MU M) with different concentrations is applied, and the drought resisting index of the Xinjiang wild apple is determined. The results show that: after 4-MU treatment, the growth status of Sinkiang Malus domestica and T337 was better than that of the non-treated group (FIGS. 2 and 3); under drought stress, the height, stem thickness, overground part dry weight, root dry weight and root-crown ratio of the Sinkiang crabapple and the T337 plant applied with 4-MU are all obviously higher than those of the unapplied group (figures 2 and 3); under the normal water supply condition, 4-MU has no obvious influence on the overground part of the rootstock (figures 2 and 3), but can obviously promote the development of the root system and promote the root-cap ratio (A, D-F in figure 2 and A, D-F in figure 3). The data show that 4-MU can improve drought resistance of Malus sieversii and T337.
2.2.2 4-MU improves relative water content of plant leaves under long-term drought stress
The relative water content of the leaves reflects the water retention capacity of the plant leaves, and the drought resistance of the plants with the leaves with higher water content under drought stress is stronger, and conversely, the drought resistance of the plants with the leaves with higher water content is weaker. The relative water content of the leaves of the Sinkiang crabapple and the T337 leaves with different concentrations of 4-MU is measured, and the result shows that: under the condition of normal water supply, the relative water content of the 4-MU leaves to the Sinkiang crabapple and the T337 leaves is not obviously different; under the stress of long-term drought, 4-MU can obviously improve the relative water content of leaves of Malus sieversii and T337 (figure 4). The data indicate that 4-MU can increase the resistance of Sinkiang Malus pumila and T337.
2.2.2 Effect of 4-MU on plant photosynthetic indices under Long-term drought stress
Under long-term moderate drought stress, stomatal opening degree is reduced, photosynthesis is weakened, and biomass accumulated by plants is reduced. Determining the photosynthetic index of the plant under different conditionsReflecting the tolerance of the plant to the external drought environment. The photosynthetic index analysis shows that: under the condition of normal water supply, photosynthetic rate (Pn), stomatal conductance (Gs) and intercellular CO of Sinkiang wild apple and T337 with different concentrations of 4-MU2Concentration and transpiration rate (Tr) were not significantly affected (fig. 5 and 6); under the stress of long-term drought, 4-MU can obviously improve the photosynthetic rate (Pn), the degree (Gs) and intercellular CO of Malus sieversii and T3372Concentration and transpiration rate (Tr) (fig. 5 and 6).
2.2.3 Effect of 4-MU on plant Water efficiency under Long-term drought stress
The water utilization efficiency is the appropriate degree for describing the growth of plants under drought stress, and is an important index for reflecting the drought resistance of the plants. The Instantaneous Water Use Efficiency (iWUE) is equal to the ratio of the photosynthetic rate to the transpiration rate. The carbon isotope technology provides powerful technical support for analyzing gas exchange and carbon absorption inside the leaves, and the water utilization efficiency of plants and the stable carbon isotope composition (delta) in dry matters of the leaves13C) The method is in positive correlation, so that the determination of the carbon isotope composition in the dry matter of the plant leaves can reflect the real water utilization state of the plant, and the method is also the most ideal index for evaluating the water utilization efficiency at present. The measurement results show that: under normal water supply conditions, 4-MU and iWUE and delta without 4-MU of Sinkiang Malus pumila and T337 are applied at different concentrations13C has no obvious difference; under drought stress, 4-MU Sinkiang Malus sieversii and T337 plants iWUE and delta were applied13The C values were all significantly higher than those of the non-applied group. The above experiments showed that 4-MU increased the water use efficiency of plants (FIG. 7).
Claims (1)
- The application of 4-methyl umbelliferone in improving the drought resistance of malus plants, wherein the improvement of the drought resistance of the plants is to promote root development, improve photosynthesis of leaves or/and improve water utilization efficiency.
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