阅读说明：本技术 一种定量植物耐低营养能力和营养利用效率的方法 (Method for quantifying low-nutrition-resistant capability and nutrition utilization efficiency of plant ) 是由 吴沿友 于睿 张承 苏跃 吴沿胜 方蕾 吴明开 王瑞 于 2019-10-10 设计创作，主要内容包括：本发明公开了一种定量植物耐低营养能力和营养利用效率的方法,属于农业工程和农作物信息检测技术领域,测定不同夹持力下植物叶片生理电阻、生理阻抗和生理电容,进一步计算植物叶片生理容抗和生理感抗；依据Nernst方程,构建植物叶片的生理电阻随夹持力变化、植物叶片的生理容抗随夹持力变化以及植物叶片的生理感抗随夹持力变化模型,利用上述三个模型的参数计算植物叶片固有生理电阻、固有生理容抗和固有生理感抗,进一步获取基于电生理参数的植物叶片营养主动转输能力和被动转输能力,最终定量出植物耐低营养能力和营养利用效率。本发明不仅可以快速、在线定量检测不同环境下不同植物耐低营养能力和营养利用效率,测定的结果具有可比性,而且还可以用生物物理指标表征不同环境下不同植物对营养的需求,为作物施肥提供科学数据。(The invention discloses a method for quantifying the low nutrition tolerance and the nutrition utilization efficiency of a plant, which belongs to the technical field of agricultural engineering and crop information detection, and is used for measuring the physiological resistance, the physiological impedance and the physiological capacitance of a plant leaf under different clamping forces and further calculating the physiological capacitance and the physiological inductance of the plant leaf; according to a Nernst equation, a model is constructed in which the physiological resistance of the plant leaves changes along with the clamping force, the physiological capacitive reactance of the plant leaves changes along with the clamping force and the physiological inductive reactance of the plant leaves changes along with the clamping force, the intrinsic physiological resistance, the intrinsic physiological capacitive reactance and the intrinsic physiological inductive reactance of the plant leaves are calculated by using the parameters of the three models, the active nutrient transferring capacity and the passive nutrient transferring capacity of the plant leaves based on electrophysiological parameters are further obtained, and finally the low nutrient tolerance capacity and the nutrient utilization efficiency of the plant are quantified. The method can rapidly and quantitatively detect the low nutrition resistance and the nutrition utilization efficiency of different plants under different environments on line, the detection result is comparable, and the biophysical indexes can be used for representing the nutrition requirements of different plants under different environments, thereby providing scientific data for crop fertilization.)

1. A method for quantifying the low nutrient tolerance and nutrient utilization efficiency of a plant, comprising the steps of:

step one, connecting a measuring device with an LCR tester;

selecting fresh branches of the plant to be detected;

collecting second unfolded leaves from the fresh branches as leaves to be detected, and soaking the leaves in distilled water;

step four, sucking water on the surface of the leaf, immediately clamping the leaf to be detected between parallel electrode plates of a detection device, setting detection voltage and frequency, setting different clamping forces by changing the mass of an iron block, and simultaneously detecting physiological capacitance, physiological resistance and physiological impedance of the plant leaf under different clamping forces in a parallel mode;

calculating physiological capacitive reactance according to the physiological capacitance of the plant leaves;

step six, calculating the physiological inductive reactance of the plant leaf according to the physiological resistance, the physiological impedance and the physiological capacitive reactance of the plant leaf;

constructing a model of physiological resistance of the plant leaves changing along with the clamping force to obtain each parameter of the model;

step eight, constructing a model of the physiological capacitive reactance of the plant leaf changing along with the clamping force to obtain each parameter of the model;

constructing a model of the physiological inductive reactance of the plant leaf along with the change of the clamping force to obtain each parameter of the model;

step ten, acquiring intrinsic physiological resistance IR of the plant leaves according to the parameters in the model in the step seven;

step eleven, acquiring inherent physiological capacitive reactance IXC of the plant leaves according to the parameters in the model in the step eight;

step twelve, acquiring inherent physiological inductive reactance IXL of the plant leaf according to the parameters in the model in the step nine;

step thirteen, calculating the inherent physiological resistance, inherent physiological capacitive reactance and reciprocal of inherent physiological inductive reactance of the plant leaves;

fourteen, taking the reciprocal of the inherent physiological resistance of the plant leaf as a reference, and obtaining the active transfer capacity NAT and the passive transfer capacity NPT of the nutrition of the plant leaf based on the electrophysiological parameters;

and step fifteen, acquiring the low nutrition tolerance RLN and the nutrition utilization efficiency NUE of the plant according to the active transfer capacity NAT and the passive transfer capacity NPT of the plant leaf nutrition based on the electrophysiological parameters.

2. The method of claim 1, wherein the method comprises the steps of: the setting method of different clamping forces in the fourth step comprises the following steps: by adding iron blocks of different masses, according to the formula of gravilogy: calculating clamping force F as (M + M) g, wherein F is the clamping force and has the unit of N; m is the mass of the iron block, and M is the mass of the plastic rod and the electrode slice, kg; g is an acceleration of gravity of 9.8N/kg.

3. The method of claim 1, wherein the method comprises the steps of: in the fifth step, a calculation formula of the physiological capacitive reactance of the plant leaves is as follows:

4. The method of claim 1, wherein the method comprises the steps of: in the sixth step, a calculation formula of the physiological inductive reactance of the plant leaf is as follows:wherein X1 is plant leaf physiological inductive reactance, Xc is plant leaf physiological capacitive reactance, Z is plant leaf physiological impedance, and R is plant leaf physiological resistance.

5. A stator according to claim 1A method for measuring the low nutrition tolerance and the nutrition utilization efficiency of plants, which is characterized in that: in the seventh step, the physiological resistance of the plant leaf changes along with the clamping force,

6. The method of claim 1, wherein the method comprises the steps of: in the step eight, the physiological capacitive reactance of the plant leaf changes along with the clamping force,

7. The method of claim 1, wherein the method comprises the steps of: in the ninth step, the physiological inductive reactance of the plant leaf changes along with the clamping force,

8. The method of claim 1, wherein the method comprises the steps of: in the step ten, the method for obtaining the intrinsic physiological resistance IR of the plant leaf according to the parameters in the model in the step seven comprises the following steps: y is IR₀+k₁(ii) a In the eleventh step, the method for obtaining the intrinsic physiological capacitive reactance IXC of the plant leaves according to the parameters in the model in the eighth step comprises the following steps: IXC ═ p₀+k₂。

9. The method of claim 1, wherein the method comprises the steps of: in the step twelve, the method for obtaining the inherent physiological inductive reactance IXL of the plant leaf according to the parameters in the model in the step nine comprises the following steps: IXL ═ q₀+k₃(ii) a Calculating the reciprocal IR of the inherent physiological resistance of the plant leaves in the step thirteen^-The calculation formula of (2):

10. the method of claim 1, wherein the method comprises the steps of: and the calculation formula of the plant leaf nutrition active transport capacity NAT based on the electrophysiological parameters in the step fourteen is as follows:the calculation formula of the plant leaf nutrition passive transport capacity NPT based on the electrophysiological parameters comprises the following steps:the calculation method of the plant low nutrition tolerance RLN in the step fifteen comprises the following steps:

Technical Field

The invention belongs to the technical field of agricultural engineering and crop information detection, and particularly relates to a method for quantifying the low-nutrient tolerance and nutrient utilization efficiency of plants, which can be used for rapidly and quantitatively detecting the substance transfer performance, the low-nutrient tolerance and the nutrient utilization efficiency of different plant leaf cell membranes in different environments on line, quantifying the low-nutrient tolerance and the nutrient utilization efficiency of different plants in the same environment on line, representing the requirements of different plants in different environments on nutrition by using biophysical indexes, and providing scientific data for crop fertilization.

Background

The cell membrane mainly comprises lipid (mainly phospholipid) (accounting for about 50% of the total amount of the cell membrane), protein (accounting for about 40% of the total amount of the cell membrane), carbohydrate (accounting for about 2% -10% of the total amount of the cell membrane) and the like; wherein the main components are protein and lipid. The phospholipid bilayer is the basic scaffold that constitutes the cell membrane. Under an electron microscope, the membrane can be divided into three layers, namely an electronic dense band (hydrophilic part) with the thickness of about 2.5nm is respectively arranged at the inner side and the outer side of the membrane, and a transparent band (hydrophobic part) with the thickness of 2.5nm is clamped in the middle.

The resistance that a cell membrane presents to a current passing through it is called the membrane resistance. Since the cell membrane is mainly composed of proteins and lipids, and thus has a large resistivity, the cell membrane becomes a major part providing the resistance of the biological tissue.

The phospholipid bilayer is the basic scaffold that constitutes the cell membrane. The membrane has hydrophilic parts near the inner and outer sides and hydrophobic part in the middle. Membrane proteins are associated with membrane lipids in two major forms: it is divided into two kinds of intrinsic protein and extrinsic protein. The intrinsic protein is directly covalently combined with the hydrophobic part of the phospholipid by the hydrophobic part, and both ends of the intrinsic protein have polarity and penetrate through the inside and the outside of the membrane; the extrinsic proteins are non-covalently bound to the outer ends of the intrinsic proteins, or to the hydrophilic heads of the phospholipid molecules. Such as carriers, specific receptors, enzymes, surface antigens. 20-30% of surface proteins (peripheral proteins) are bonded to lipids on both sides of the membrane with charged amino acids or groups, polar groups; 70-80% of the binding proteins (intrinsic proteins) are bound to lipid molecules through one or more hydrophobic alpha-helices (formed by 20-30 hydrophobic amino acids absorbed, 3.6 amino acid residues per turn, corresponding to the membrane thickness. adjacent alpha-helices are linked by linear peptides on the inside and outside of the membrane), i.e., hydrophobic hydroxyl groups in the membrane. Such a cell membrane structure results in its capacitive and inductive properties. The type and amount of surface proteins (peripheral proteins) among others determines the magnitude of their capacitance, and the type and amount of binding proteins (intrinsic proteins), especially of transport proteins among others, determines the magnitude of their inductance.

There are two main classes of transporters present on the cell membrane, namely: carrier proteins (carrier proteins) and channel proteins (channel proteins). Carrier proteins, also known as carriers (carriers), permeases (permeases) and transporters (transporters), are capable of binding to a specific solute and, by a change in their conformation, of transferring the solute bound to it to the other side of the membrane, and some of them require energy to drive, such as: various ATP-driven ion pumps; some do not require energy to transport materials in a freely diffusing manner, such as: valinase. The channel proteins bind weakly to the transported substance, form hydrophilic channels that allow specific solutes to pass through when the channels are open, and all channel proteins transport solutes in a freely diffusing manner.

The cell membrane is a barrier for preventing extracellular substances from freely entering cells, and ensures the relative stability of the intracellular environment, so that various biochemical reactions can be orderly operated. However, the cells must exchange information, substances and energy with the surrounding environment to perform a specific physiological function, and therefore, the cells must have a substance transport system for obtaining desired substances and discharging metabolic wastes. It is estimated that the proteins on the cell membrane involved in substance transport account for 15-30% of the proteins encoded by nuclear genes, and the energy used by the cells in substance transport amounts to two thirds of the total energy consumed by the cells. From this, it can also be seen that the substance transport ability of the cell is determined by the kind and amount of the surface protein and the binding protein in the cell membrane.

The components and structure of the cell membrane play an important role in the operation of substances, and the different components and structures of the cell membrane determine the electrophysiological characteristics of cells and constituent organs thereof and also determine the absorption and transport functions of different nutrients. The ratio of phospholipids, surface proteins (peripheral proteins) and binding proteins (intrinsic proteins) on cell membranes strongly influences the transport capacity of cell material and influences the metabolism of inorganic nutrients, which are closely related to the utilization efficiency of nutrient elements and ultimately influence the utilization efficiency of plant nutrients. In addition, the proportion of the binding protein (intrinsic protein) is closely related to the active transport of some nutrient elements, the proportion of the cell substance transport capacity caused by the binding protein to the total substance transport capacity determines the active transport capacity of the nutrient elements, and the strength of the active transport capacity of the nutrient elements is closely related to the low nutrition tolerance of plants. Therefore, in order to determine the contribution of phospholipids, surface proteins (peripheral proteins) and binding proteins (intrinsic proteins) on cell membranes to the operation of cell membrane substances and the proportion of the active transport capacity of nutrient elements to the total transport capacity of the substances, the invention takes plant leaves as an investigation organ, jointly deduces the physiological resistance of the plant leaves along with the change of clamping force, the physiological volume resistance of the plant leaves along with the change of clamping force and the physiological inductance of the plant leaves along with the change of clamping force according to an nernst equation, calculates the inherent physiological resistance, the inherent physiological volume resistance and the inherent physiological inductance of the plant leaves by using the parameters of the three models, further obtains the active transport capacity and the passive transport capacity of the plant leaf nutrition based on electrophysiological parameters, and finally quantifies the low nutrition-resistant capacity and the nutrition utilization efficiency of the plant. The method can rapidly and quantitatively detect the low-nutrition-resistant capability and the nutrition utilization efficiency of different plants under different environments on line, the detection result is comparable, and the biophysical indexes can be used for representing the requirements of different plants under different environments on nutrients and the transport capability of different plant metabolites under different environments, thereby providing scientific data for crop fertilization.

Disclosure of Invention

The invention aims to provide a method for quantifying the low nutrition tolerance and the nutrition utilization efficiency of plants, which not only fills the blank that the biophysical indexes are used for representing the active nutrition transfer capacity and the passive nutrition transfer capacity, but also fills the blank that the biophysical indexes are used for representing the requirements of different plants on nutrients in different environments and the transport capacities of different plant metabolites in different environments, and provides a scientific basis for accurate fertilization of crops.

In order to solve the technical problems, the invention adopts the following specific technical scheme:

a method of quantifying the low nutrient tolerance and nutrient use efficiency of a plant comprising the steps of:

step one, connecting a measuring device with an LCR tester;

selecting a fresh branch of the plant to be detected, and wrapping the base of the branch;

collecting second unfolded leaves from the fresh branches as leaves to be detected, and soaking the leaves in distilled water for 30 minutes;

step four, sucking water on the surface of the leaf, immediately clamping the leaf to be detected between parallel electrode plates of a detection device, setting detection voltage and frequency, setting different clamping forces by changing the mass of an iron block, and simultaneously detecting physiological capacitance, physiological resistance and physiological impedance of the plant leaf under different clamping forces in a parallel mode;

calculating physiological capacitive reactance according to the physiological capacitance of the plant leaves;

step six, calculating the physiological inductive reactance of the plant leaf according to the physiological resistance, the physiological impedance and the physiological capacitive reactance of the plant leaf;

constructing a model of physiological resistance of the plant leaves changing along with the clamping force to obtain each parameter of the model;

step eight, constructing a model of the physiological capacitive reactance of the plant leaf changing along with the clamping force to obtain each parameter of the model;

constructing a model of the physiological inductive reactance of the plant leaf along with the change of the clamping force to obtain each parameter of the model;

step ten, acquiring intrinsic physiological resistance IR of the plant leaves according to the parameters in the model in the step seven;

step eleven, acquiring inherent physiological capacitive reactance IXC of the plant leaves according to the parameters in the model in the step eight;

step twelve, acquiring inherent physiological inductive reactance IXL of the plant leaf according to the parameters in the model in the step nine;

step thirteen, calculating the inherent physiological resistance, inherent physiological capacitive reactance and reciprocal of inherent physiological inductive reactance of the plant leaves;

fourteen, taking the reciprocal of the inherent physiological resistance of the plant leaf as a reference, and obtaining the active transfer capacity NAT and the passive transfer capacity NPT of the nutrition of the plant leaf based on the electrophysiological parameters;

and step fifteen, acquiring the low nutrition tolerance RLN and the nutrition utilization efficiency NUE of the plant according to the active transfer capacity NAT and the passive transfer capacity NPT of the plant leaf nutrition based on the electrophysiological parameters.

Furthermore, the measuring device in the first step comprises a support (1), foam plates (2), electrode plates (3), leads (4), iron blocks (5), a plastic rod (6) and a fixing clamp (7), wherein the support (1) is of a rectangular frame structure, one side of the support is open, a through hole is formed in the upper end of the support (1) and used for the plastic rod (6) to extend into, the inward side of the lower end of the support (1) and the bottom end of the plastic rod (6) are respectively adhered with the two foam plates (2), the electrode plates (3) are embedded in the foam plates (2), the leads (4) are respectively led out from the two electrode plates (3), the iron blocks (5) with fixed quality can be placed on the foam plates (2) of the plastic rod (6), and one end, located inside the support, of the plastic rod (6) is fixed by the fixing clamp (7); the electrode plate (3) is a circular electrode plate, and the electrode plate (3) is made of copper.

Further, the second unfolded leaf on the fresh branch in the third step is based on the principle from top to bottom, and the leaf which is just completely developed and is completely unfolded on the fresh branch is taken as the first completely unfolded leaf, and so on; respectively, the second fully expanded leaf, the third fully expanded leaf, etc.

Further, the setting method of the different clamping forces in the fourth step is as follows: by adding iron blocks of different masses, according to the formula of gravilogy: calculating clamping force F as (M + M) g, wherein F is the clamping force and has the unit of N; m is the mass of the iron block, and M is the mass of the plastic rod and the electrode slice, kg; g is an acceleration of gravity of 9.8N/kg.

Further, the calculation formula of the physiological capacitive reactance of the plant leaves in the fifth step is as follows:

wherein Xc is the physiological capacitive reactance of the plant leaves, C is the physiological capacitance of the plant leaves, f is the test frequency, and pi is the circumference ratio equal to 3.1416.

Further, a calculation formula of the physiological inductive reactance of the plant leaf is as follows:

wherein X1 is plant leaf physiological inductive reactance, Xc is plant leaf physiological capacitive reactance, Z is plant leaf physiological impedance, and R is plant leaf physiological resistance.

Further, in the seventh step, the physiological resistance of the plant leaf changes along with the clamping force,

the model is based on the Nernst equation

Deduced, wherein R is a physiological resistance, E is an electromotive force, and E⁰Is a standard electromotive force, R₀Is an ideal gas constant, T is temperature, C_iConcentration of dielectric substances, C, in response to physiological resistance in cell membranes_oConcentration of dielectric substances in response to physiological resistance outside cell membrane, f₀Concentration C of dielectric substance responsive to physiological resistance in cell membrane_iProportional coefficient converted from physiological resistance, total dielectric substance C of intra-membrane and extra-membrane response physiological resistance_T＝C_i+C_o，F₀Is the Faraday constant, n_RIs the number of dielectric mass transfers in response to physiological resistance; e can be used for doing work, PV is proportional to PV and is a E, a is the coefficient of converting electromotive force into metabolic energy, v is the volume of plant cells, P is the pressure to which the plant cells are subjected, and the pressure P is expressed by the pressure formula

Calculating F as clamping force, S as effective area under the action of the polar plate and d as specific effective thickness of the plant leaf;the deformation is as follows:

and is further transformed into

Due to the specific effective thickness of the plant leaves

Therefore, the temperature of the molten metal is controlled,

the deformation is as follows:order to

The model of the physiological resistance of the plant leaf changing along with the clamping force can be deformed into

Wherein y is₀、k₁And b₁Are parameters of the model.

Further, in the eighth step, the physiological capacitive reactance of the plant leaf is clamped along with the clampingA model of the change in force is generated,

the model is based on the Nernst equation

Deduced, wherein Xc is physiological capacitive reactance, E is electromotive force, E⁰Is a standard electromotive force, R₀Is the ideal gas constant, T is the temperature, Q_iDielectric substance concentration, Q, in response to physiological capacitive reactance in cellular membranes_oConcentration of dielectric substances in response to physiological capacitive impedance outside cell membrane, J₀Dielectric substance concentration Q that is responsive to physiological capacitive reactance in cellular membranes_iThe ratio coefficient of conversion between the dielectric substance and the physiological capacitive reactance, and the total amount of the dielectric substance Q which responds to the physiological capacitive reactance inside and outside the membrane is Q_i+Q_o，F₀Is the Faraday constant, n_XCIs the number of dielectric mass transfers in response to physiological capacitive reactance; e can be used for doing work, PV is proportional to PV and is a E, a is the coefficient of converting electromotive force into metabolic energy, V is the volume of plant cells, P is the pressure to which the plant cells are subjected, and the pressure P is expressed by the pressure formula

Calculating F as clamping force, s as effective area under the action of the polar plate, and d as specific effective thickness of the plant leaf;

the deformation is as follows:

and is further transformed into

Due to the specific effective thickness of the plant leaves

Therefore, the temperature of the molten metal is controlled, the deformation is as follows:order to

The model of the physiological capacitance of the plant leaf changing along with the clamping force can be deformed into

Wherein p is₀、k₂And b₂Are parameters of the model.

Further, in the ninth step, the physiological inductive reactance of the plant leaf is changed along with the clamping force,

the model is based on the Nernst equation

Deduced, wherein X1 is a physiological inductive reactance, E is an electromotive force, E⁰Is a standard electromotive force, R₀Is the ideal gas constant, T is the temperature, M_iDielectric concentration, M, in response to physiological inductance within the cell membrane_oConcentration of dielectric substances, L, in response to physiological inductance outside the cell membrane₀Dielectric substance concentration M being responsive to physiological inductive reactance in cell membrane_iThe proportionality coefficient for conversion between the dielectric substance and physiological inductive reactance, and the total dielectric substance M of the intra-membrane and extra-membrane response physiological inductive reactance_T＝M_i+M_o，F₀Is the Faraday constant, n_xLIs the number of dielectric mass transfers in response to physiological inductance; e can be used for doing work, PV is proportional to PV a E, a is the coefficient of converting electromotive force into metabolic energy, and V is plant finenessCell volume, P is the pressure to which the plant cells are subjected, and the pressure P is expressed by the pressure equation

Calculating F as clamping force, s as effective area under the action of the polar plate, and d as specific effective thickness of the plant leaf;

the deformation is as follows:

and is further transformed into

Due to the specific effective thickness of the plant leavesTherefore, the temperature of the molten metal is controlled,

the deformation is as follows:

order toThe model of the physiological inductive reactance of the plant leaf changing along with the clamping force can be deformed into

Wherein q is₀、k₃And b₃Are parameters of the model.

Further, in the step ten, the method for obtaining the intrinsic physiological resistance IR of the plant leaf according to the parameters in the model in the step seven comprises the following steps: y is IR₀+k₁。

Further, in the eleventh step, the inherent physiology of the plant leaf is obtained according to the parameters in the model in the eighth stepThe capacitive reactance IXC method comprises the following steps: IXC ═ p₀+k₂。

Further, in the twelfth step, the method for obtaining the inherent physiological inductive reactance IXL of the plant leaf according to the parameters in the ninth model comprises the following steps: IXL ═ q₀+k₃。

Further, the calculation formula for calculating the reciprocal of the intrinsic physiological resistance IR-of the plant leaf in the step thirteen is as follows:

the calculation formula of the inherent physiological capacitive reactance reciprocal IXC of the plant leaves is as follows:

the calculation formula of the reciprocal of the inherent physiological inductive reactance IXL-of the plant leaf is as follows:

further, in the fourteenth step, the calculation formula of the active plant leaf nutrition transferring capability NAT based on electrophysiological parameters is as follows:

the calculation formula of the plant leaf nutrition passive transport capacity NPT based on the electrophysiological parameters comprises the following steps:

further, the calculation method of the plant low-nutrition-tolerance RLN in the step fifteen is as follows:

unit%; the method for calculating the NUE of the plant nutrition utilization efficiency comprises the following steps:and has no unit.

The invention has the following beneficial effects:

1. the invention can rapidly and quantitatively detect the inherent nutrition active transfusion capability and passive transfusion capability of different plants under different environments on line, and the detection result is not changed due to the change of the detection conditions and has comparability.

2. The invention represents the requirements of different plants on nutrients by electrophysiological indexes through quantitative determination of the low nutrition tolerance and the nutrition utilization efficiency of the plants.

3. The invention uses electrophysiological indexes to represent the inherent transport capacity of different plant metabolites under different environments by measuring the contribution of phospholipids, surface proteins (peripheral proteins) and binding proteins (intrinsic proteins) on cell membranes to the operation of cell membrane substances, provides quantitative data for comparing the change of cell membrane functions of different plants under different environments, and provides scientific basis for accurate fertilization of crops.

4. The invention is simple and convenient, has wide applicability and low price of required instruments.

Drawings

FIG. 1 is a structural model of a cell membrane;

FIG. 2 is a schematic view showing the structure of the measuring apparatus of the present invention;

in the figure: 1. a support; 2. a foam board; 3. an electrode plate; 4. an electrical lead; 5. an iron block; 6. a plastic rod; 7. and (4) fixing clips.

Detailed Description

The invention is further described with reference to the following figures and examples.

The basic principle of the invention is as follows:

from the formula of gravimetry:

F＝(M+m)g (1)

wherein F is gravity (clamping force), N; m is the mass of the iron block, and M is the mass of the plastic rod and the electrode slice, kg; g is the acceleration of gravity of 9.8, N/kg.

The cytosol in the leaf is used as a dielectric medium, and the leaf is clamped between two parallel plate capacitor plates of a parallel plate capacitor to form the parallel plate capacitance sensor. The physiological capacitance of the plant leaf under different clamping forces is obtained by adding iron blocks with certain mass, and different pressures can lead to different changes of the concentration of the cytosol in the leaf, so that the elasticity and plasticity of leaf cells are changed, the change of the cytosol dielectric constant of leaf tissue between two capacitor plates is caused, and the electrophysiological indexes of plant physiological capacitance, resistance, impedance and the like are influenced.

The calculation formula of the physiological capacitive reactance of the plant leaves is as follows:

wherein Xc is the physiological capacitive reactance of the plant leaves, C is the physiological capacitance of the plant leaves, f is the test frequency, and pi is the circumference ratio equal to 3.1416.

The physiological resistance, physiological impedance and physiological capacitance of the plant leaves are measured in a parallel mode; therefore, the calculation formula of the physiological inductive reactance of the plant leaf is as follows:

wherein X1 is plant leaf physiological inductive reactance, Xc is plant leaf physiological capacitive reactance, z is plant leaf physiological impedance, and R is plant leaf physiological resistance.

Since the resistive current is caused by the dielectric substance, it is determined by factors such as the degree of permeability of the film to various dielectric substances and the presence or absence of a large amount of the dielectric substance. The external excitation changes the permeability of the dielectric substance, the concentration of the internal and external dielectric substances is influenced, the concentration difference of the internal and external dielectric substances obeys the Nemst equation, the physiological resistance is inversely proportional to the conductivity, and the conductivity is proportional to the concentration of the dielectric substance in the cell, so that the relationship between the physiological resistance of the cell and the external excitation can be deduced.

The water content of plant cells is related to the elasticity of plant leaf cells, and under different clamping forces, the permeability of different plant cell membranes is changed differently, so that the physiological resistance of the plant cell membranes is different.

The expression of the nernst equation is as follows (2):

wherein E is electromotive force; e⁰Is a standard electromotive force; r₀Is an ideal gas constant equal to 8.314570J.K^-1.mol^-1T is temperature, singlyA bit K; c_iConcentration of dielectric substances, C, in response to physiological resistance in cell membranes_oThe concentration of the dielectric substance responding to the physiological resistance outside the cell membrane and the total amount C of the dielectric substance responding to the physiological resistance inside and outside the cell membrane_T＝C_i+C_o，F₀Is the Faraday constant, equal to 96485C.mol^-1(ii) a nR is the number of dielectric mass transfers in mol in response to physiological resistance.

The internal energy of electromotive force E can be converted into pressure to do work, and PV is proportional to PV ═ aE, namely:

wherein: p is the pressure applied to the plant cell, a is the electromotive force conversion energy coefficient, and v is the plant cell volume;

the pressure P to which the plant cells are subjected can be determined by a pressure formula:wherein F is the clamping force, and s is the effective area under the action of the polar plate;

in mesophyllic cells, the vacuole and the cytoplasm occupy the vast majority of the intracellular space. For mesophyllic cells, C_oAnd C_iThe sum is constant and equal to the total amount C of dielectric substances responding to physiological resistance inside and outside the membrane_T，C_iIt is proportional to the conductivity, which is the inverse of the resistance R, and, therefore,can be expressed as

Wherein R is resistance, f₀Concentration C of dielectric substance responsive to physiological resistance in cell membrane_iAnd the proportionality coefficient of the conversion between resistance, therefore, (3) can become:

(4) is transformed to obtain

(5) Is transformed to obtain

(6) The two sides of the formula are taken as indexes and can be changed into:

further modified, it is possible to obtain:

r in the formula (8) is a physiological resistance due to the specific effective thickness of the plant leaves(8) The formula can be deformed into:

d, a and E in formula (9) for the same blade to be tested in the same environment⁰、R₀、T、n_R、F₀、C_T、f₀Are all constant values; order to

Therefore, equation (9) can be transformed into:

(10) in the formula y₀、k₁And b₁Are parameters of the model. When F is 0 and is substituted into the formula (10), the plant leaf intrinsic physiological resistance IR is obtained: y is IR₀+k₁。

In the capacitive reactance measurement of the same object under the same environment, the capacitive reactance mainly depends on the concentration of dielectric substances responding to physiological capacitive reactance inside and outside the membrane, so the permeability of the membrane to various dielectric substances responding to physiological capacitive reactance determines the size of the cellular capacitive reactance, and for the leaf, the capacitive reactance further depends on the concentration of the dielectric substances responding to physiological capacitive reactance inside and outside the membrane. The external excitation changes the membrane permeability of the dielectric substance, the concentration of the dielectric substance responding to the physiological capacitive reactance inside and outside the membrane is influenced, the concentration difference of the dielectric substance responding to the physiological capacitive reactance inside and outside the membrane also obeys a Nemst (Nemst) equation, and when the concentration of the dielectric substance responding to the physiological capacitive reactance outside the membrane is constant, the physiological capacitive reactance is inversely proportional to the concentration of the dielectric substance responding to the physiological capacitive reactance inside the cell, so that the relation between the physiological capacitive reactance inside the cell and the external excitation can be deduced.

The water content of plant cell is related to the elasticity of plant leaf cell, and under different clamping forces, the permeability of dielectric substance responding to physiological capacitive reactance of different plant cell membranes is changed differently, so that the physiological capacitive reactance is different.

The expression of the nernst equation is as shown in equation (11):

wherein E is electromotive force, E⁰Is a standard electromotive force, R₀Is an ideal gas constant equal to 8.314570J.K^-1.mol^-1(ii) a T is temperature, in K; q_iDielectric substance concentration, Q, in response to physiological capacitive reactance in cellular membranes_oThe total amount of dielectric substance Q ═ Q for responding to physiological capacitive reactance outside cell membrane and for responding to physiological capacitive reactance outside membrane_i+Q_o，F₀Is the Faraday constant, equal to 96485C.mol^-1；n_XCIs the number of dielectric material transitions in mol in response to physiological capacitive reactance.

The internal energy of electromotive force E can be converted into pressure to do work, and PV is proportional to PV ═ aE, namely:

wherein: p is the pressure to which the plant cell is subjected, a is the electromotive force conversion energy coefficient, and V is the plant cell volume;

the pressure P to which the plant cells are subjected can be determined by a pressure formula:

wherein F is the clamping force and s is the effective area under the action of the polar plate;

in mesophyllic cells, the vacuole and the cytoplasm occupy the vast majority of the intracellular space. For mesophyll cells, Q_oAnd Q_iThe sum is certain and equal to the total amount of dielectric substances Q, Q responding to physiological capacitive reactance inside and outside the membrane_iIs proportional to the conductivity of the dielectric material responsive to the physiological capacitive impedance, which is the inverse of the capacitive impedance Xc, and, therefore,can be expressed as

Xc is a capacitive impedance, J0 is the proportionality coefficient for the transformation between the dielectric substance concentration Qi and the capacitive impedance in response to physiological capacitive impedance within the cell membrane, and thus (12) can become:

(13) is transformed to obtain

(14) Can become:

(15) the two sides of the formula are taken as indexes and can be changed into:

further modified, it is possible to obtain:

xc in formula (17) is a physiological capacitive reactance due to the specific effective thickness of the plant leaves

(17) The formula can be deformed into:

for the same blade to be measured in the same environment, (18) formula (d, a, E)⁰、R₀、T、n_XC、F₀、Q、J₀Are all constant values, order

Thus, equation (18) can be transformed as:

(19) in the formula p₀、k₂And b₂Are parameters of the model. When F is 0, substituting into formula (19), the plant leaf intrinsic physiological capacitive reactance IXC is obtained: IXC ═ p₀+k₂。

Similarly, the permeability of the dielectric substance responding to physiological inductance of different plant cell membranes is changed differently under different clamping forces, so that the physiological inductance is different.

The expression of the nernst equation is as in equation (20):

wherein E is electromotive force, E⁰Is a standard electromotive force, R₀Is an ideal gas constant equal to 8.314570J.K^-1.mol^-1(ii) a T is temperature, in K; m_iDielectric concentration, M, in response to physiological inductance within the cell membrane_oThe total amount of dielectric substance M is the concentration of dielectric substance responding to physiological inductance outside the cell membrane_T＝M_i+M_o，F₀Is the Faraday constant, equal to 96485C.mol^-1；n_XLIs the number of dielectric material transfers in mol in response to physiological inductance.

The internal energy of electromotive force E can be converted into pressure to do work, and PV is proportional to PV ═ aE, namely:

wherein: p is the pressure to which the plant cell is subjected, a is the electromotive force conversion energy coefficient, and V is the plant cell volume;

the pressure P to which the plant cells are subjected can be determined by a pressure formula:

wherein F is the clamping force and s is the effective area under the action of the polar plate;

in mesophyllic cells, the vacuole and the cytoplasm occupy the vast majority of the intracellular space. For mesophyllic cells, M_oAnd M_iThe sum is a certain amount, which is equal to the total amount M of dielectric substances responding to physiological inductive reactance inside and outside the membrane_T，M_iIt is proportional to the conductivity of the dielectric material in response to the physiological impedance, which is the reciprocal of the impedance X1, and therefore,

can be expressed as

X1 is inductive reactance, L₀Is the physiological sensation of response in cell membraneThe proportionality coefficient for the conversion between the dielectric concentration Mi of the reactance and the inductive reactance, and therefore, the formula (21) can be changed as follows:

(22) is transformed to obtain

(24) Can become:

(24) the two sides of the formula are taken as indexes and can be changed into:

further modified, it is possible to obtain:

x1 in formula (26) is physiological inductive reactance due to the specific effective thickness of the plant leaves

(26) The formula can be deformed into:

for the same blade to be measured in the same environment, (27) formula (d, a, E)⁰、R₀、T、n_XL、F₀、M_T、L₀Are all constant values, order

Therefore, equation (27) can be transformed into:

(28) in the formula q₀、k₃And b₃Are parameters of the model. When F is substituted into 0 in formula (28), the plant leaf intrinsic physiological inductive resistance IXL is obtained: IXL ═ q₀+k₃。

Calculation formula of intrinsic physiological resistance IR of plant:

wherein IR₁、IR₂、IR₃、…IR_nAssuming that the intrinsic resistance of each unit cell membrane is equal, i.e., IR₁＝IR₂＝IR₃＝…＝IR_n＝IR₀Then, the calculation formula of the intrinsic physiological resistance of the plant is:where n can then be characterized as the amount of proteins and lipids that cause the electrical resistance of the biological tissue.

Calculation formula of inherent physiological capacitive reactance IXC of plant:

wherein IXC₁、IXC₂、IXC₃、…IXC_pAssuming the inherent capacitive reactance of each cell membrane unit is equal, i.e. IXC₁＝IXC₂＝IXC₃＝…＝IXC_p＝IXC₀Then, the calculation formula of the inherent physiological capacitive reactance of the plant is as follows:

where p can then be characterized as the number of proteins, in particular surface proteins (peripheral proteins), which cause capacitive resistance in biological tissues.

Calculation formula of inherent physiological inductive reactance IXL of plant:wherein IXL₁、IXL₂、IXL₃、…IXL_qThe inherent inductance of each cell membrane unit is assumed to be equal, i.e. IXL₁＝IXL₂＝IXL₃＝…＝IXL_q＝IXL₀Then, the calculation formula of the intrinsic physiological impedance of the plant is:

wherein q can then be characterized by the number of protein-binding proteins (intrinsic proteins) which cause an inductive resistance in biological tissues, in particular transport proteins therein.

Inherent physiological inductive reactance inverse IXL of plant leaf^-The calculation formula of (2):reciprocal IXC of inherent physiological capacitive reactance of plant leaves^-The calculation formula of (2):

the calculation formula of the reciprocal R-of the inherent physiological resistance of the plant leaf is as follows:the ratio of cellular material transport capacity due to surface proteins (peripheral proteins) to total material transport capacity determines the passive transport capacity of the nutrient elements, and the ratio of cellular material transport capacity due to binding proteins to total material transport capacity determines the active transport capacity of the nutrient elements. Because, the active nutrient transferring capacity of the plant leaves based on the electrophysiological parameters

Simultaneously due to the same plant

To a certain extent, NAT can therefore be characterized as the active transport capacity of plant nutrients. Plant leaf nutrition passive transfusion capability based on electrophysiological parameters

Due to the same plant

To a certain extent, NPT can therefore be characterised as causing the passive transport of plant nutrients. The plant active transport capacity determines the minimum ion absorption concentration and thus the plant low-nutrition resistance, so that the plant low-nutrition resistance can be the ratio of the plant active transport capacity to the total plant nutrition transport capacity. The total transport capacity of the plant nutrition is NAT + NPT, so that the plant can tolerate low nutrition

Unit%; the plant nutrient utilization efficiency is expressed as

And has no unit.

A device for measuring the low nutrition tolerance and the nutrition utilization efficiency of plants comprises a bracket 1, a foam plate 2, an electrode plate 3, an electric lead 4, an iron block 5, a plastic rod 6 and a fixing clamp 7, as shown in figure 2; the bracket 1 is of a rectangular frame structure, one side of the bracket is open, the upper end of the bracket 1 is provided with a through hole for a plastic rod 6 to extend into, the inward side of the lower end of the bracket 1 and the bottom end of the plastic rod 6 are respectively adhered with two foam plates 2, electrode plates 3 are embedded in the foam plates 2, a lead 4 is respectively led out from each of the two electrode plates 3 and is used for being connected with an LCR tester (HIOKI 3532-50 type, Japan day place), an iron block 5 with fixed mass can be placed on the foam plates 2 of the plastic rod 6, and the physiological resistance, the physiological impedance and the physiological capacitance of the plant leaves are measured in a parallel connection mode; one end of the plastic rod 6, which is positioned in the bracket, is fixed by a fixing clamp 7, and when the lower end of the plastic rod is combined with the end of the bracket, the two electrode plates 3 are completely and correspondingly combined together; the electrode plate 3 is a circular electrode plate made of copper to reduce the edge effect of the electrode.

The method comprises the following steps: when the device is used, two wires 4 of the device are connected with a 9140 four-terminal test probe of an LCR tester, then the plastic rod 6 is lifted, two electrode plates 3 clamp plant leaves to be measured, the diameter of each electrode plate is 10mm, the measurement voltage is set to be 1.5V, the measurement frequency is 3000Hz, the mass of the plastic rod and the electrode plates and the mass of the iron block 5 are calibrated, and the physiological resistance, the physiological impedance and the physiological capacitance of the plant leaves under different clamping forces are measured in a parallel mode.