CN108872317B - Method for measuring dielectric substance transfer number of plant leaf cells - Google Patents
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Abstract
The invention discloses a method for measuring the transfer number of dielectric substances of plant leaf cells, which belongs to the technical field of biophysical information detection.A polar plate is connected with an LCR tester through a lead when a measuring device is used, the two polar plates clamp plant leaves to be measured, the physiological resistance, the physiological impedance and the physiological capacitance of the plant leaves under different clamping forces are measured simultaneously in a parallel mode, and the physiological capacitive reactance and the physiological inductive reactance of the plant leaves are further calculated; according to a Nernst equation, a model is constructed in which the physiological resistance of the plant leaves changes with the clamping force, the physiological capacitance of the plant leaves changes with the clamping force and the physiological inductance of the plant leaves changes with the clamping force, and the transfer percentages of the dielectric substances of different types are calculated by utilizing all parameters of the three models in a combined mode. The invention not only can rapidly and quantitatively detect the transfer percentages of various dielectric substances of different types of different plant leaves under different environments on line, but also can represent the communication characteristics of the water and the substances of different plant leaves in the system under different environments by using biophysical indexes.
Description
Technical Field
The invention belongs to the technical field of biophysical information detection, and particularly relates to a method for determining the transfer number of dielectric substances of plant leaf cells, which can quickly detect the communication characteristics of water and substances of plant leaves in a system and provide data support for the explanation of the substance transportation characteristics of plant leaf cell membranes.
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.
As shown in fig. 1, 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 membrane proteins are bound to the membrane lipids in two forms, an intrinsic protein and an extrinsic protein, the intrinsic protein being covalently bound with hydrophobic moieties directly to the phospholipid, both ends being polar, penetrating the inside and outside of the membrane, the extrinsic protein being non-covalently bound to the outer ends of the intrinsic protein or to the hydrophilic heads of the phospholipid molecules, for example carriers, specific receptors, enzymes, surface antigens, 20% to 30% of the surface proteins (peripherins) being bound with charged amino acids or groups, polar groups, to the lipids on both sides of the membrane, 70% to 80% of the bound proteins (intrinsic proteins) being formed by absorption of one or several hydrophobic α -helices (20-30 hydrophobic amino acids, 3.6 amino acid residues per turn, corresponding to the membrane thickness, adjacent α -helices being linked with linear peptides on both inside and outside of the membrane, i.e. hydrophobic hydroxyl groups in the membrane, are bound to the lipid molecules.
There are two main classes of transporters present on the cell membrane, namely: carrier proteins (carrierprotein) and channel proteins (channel protein). 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 APT 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 transport capacity of plant leaf cells is related to various physiological activities of water metabolism, transport of photosynthetic products, nitrate reduction and the like of plants. In order to determine the contribution of phospholipids, surface proteins (peripheral proteins) and binding proteins (intrinsic proteins) on cell membranes to the movement of cell membrane substances, the plant leaf is taken as an investigation organ, a model that the physiological resistance of the plant leaf changes along with the clamping force, the physiological capacitive resistance of the plant leaf changes along with the clamping force and the physiological inductive resistance of the plant leaf changes along with the clamping force is jointly deduced according to the Nernst equation, and the transfer percentages of different types of dielectric substances are jointly calculated by utilizing the parameters of the three models. The method not only can rapidly and quantitatively detect the transfer percentages of various dielectric substances of different types of different plant leaves under different environments on line, and the determination result has comparability, but also can represent the communication characteristics of water and substances of different plant leaves in systems under different environments by using biophysical indexes, determine the contribution parts of phospholipid, surface protein and binding protein on cell membranes to the operation of cell membrane substances, and provide scientific data for clarifying complex biological rules and source-base relations of plant organs.
Disclosure of Invention
The invention aims to provide a method for measuring the dielectric substance transfer number of leaf cells of plants, fills the blank that the transfer capacity of different substances of the leaf cells is represented by biophysical indexes, and provides a mode for quantifying the composition structure and the function of plant leaf cell membranes.
In order to solve the technical problems, the invention adopts the following specific technical scheme:
a method for determining the dielectric substance transfer number of plant leaf cells comprises the following steps:
step one, connecting a measuring device with an LCR tester;
selecting a fresh branch of a plant to be detected, wrapping the base of the branch, and bringing the branch to a laboratory;
collecting leaves to be detected from the fresh branches, 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, respectively obtaining the dielectric substance transfer number Kn of the k-type response physiological resistance according to the parameters in the model in the step sevenRAnd b-type dielectric material transfer number Bn responding to physiological resistanceR;
Step eleven, respectively obtaining the dielectric substance transfer number Kn of the k-type response physiological capacitive reactance according to the parameters in the model in the step eightXCAnd b-type dielectric substance transfer number Bn in response to physiological capacitive impedanceXC;
Step twelve, respectively obtaining the dielectric substance transfer number Kn of the k-type response physiological inductive reactance according to the parameters in the model in the step nineXLAnd b-type dielectric material transfer number Bn responding to physiological inductive reactanceXL;
Thirteen, responding to the dielectric substance transfer number Kn of the physiological resistance according to the k typeRK-type dielectric substance transfer number Kn of response physiological capacitive reactanceXCAnd k-type dielectric substance transfer number Kn in response to physiological inductive reactanceXLObtaining the k-type total dielectric material transfer number KnT;
Fourteen, responding to the dielectric material transfer number Bn of the physiological resistance according to type bRType b dielectric material transfer number Bn responding to physiological capacitive impedanceXCAnd b-type dielectric material transfer number Bn responding to physiological inductive reactanceXLObtaining the b-type total dielectric material transfer number BnT;
Step fifteen, according to KnR、KnXC、KnXLAnd KnTObtaining KPn dielectric mass transfer percentages of k-type response physiological resistanceRDielectric mass transfer percentage of k-type response to physiological capacitive reactance KPnXCAnd k-type response physiological inductive reactancePercentage of dielectric material transfer KPnXL;
Step sixteen, according to BnR、BnXC、BnXLAnd BnTObtaining the dielectric substance transfer percentage BPn of the b-type response physiological resistance respectivelyRType b dielectric mass transfer percentage BPn in response to physiological capacitive impedanceXCAnd b-type dielectric mass transfer percentage BPn in response to physiological inductanceXL。
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 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, in the fifth step, a calculation formula of the physiological capacitive reactance of the plant leaf 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 Xl is plant leaf physiological inductive reactance, Xc is plant leaf physiological capacitive reactance, and Z is plant leafPhysiological impedance, R is the physiological resistance of the plant leaf.
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 equationDeduced, 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, CiConcentration of dielectric substances, C, in response to physiological resistance in cell membranesoConcentration of dielectric substances in response to physiological resistance outside cell membrane, f0Concentration C of dielectric substance responsive to physiological resistance in cell membraneiProportional coefficient converted from physiological resistance, total dielectric substance C of intra-membrane and extra-membrane response physiological resistanceT=Ci+Co,F0Is the Faraday constant, nRIs the number of dielectric mass transfers in response to physiological resistance; e can be used for doing work, PV is proportional to PV (a is aE), a is the coefficient of converting electromotive force into metabolic energy, V is the volume of the plant cells, P is the pressure to which the plant cells are subjected, and the pressure P is expressed by a pressure formulaCalculating 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 intoDue 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 resistance of the plant leaf changing along with the clamping force can be deformed intoWherein y is0、k1And b1Are parameters of the model.
Further, in the eighth step, the physiological volume resistance of the plant leaf changes along with the clamping force, the model is based on the Nernst equationDeduced, wherein Xc is physiological capacitive reactance, E is electromotive force, E0Is a standard electromotive force, R0Is the ideal gas constant, T is the temperature, QiDielectric substance concentration, Q, in response to physiological capacitive reactance in cellular membranesoConcentration 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 membranesiThe 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 Qi+Qo,F0Is the Faraday constant, nXCIs the number of dielectric mass transfers in response to physiological capacitive reactance; e can be used for doing work, PV is proportional to PV (a is aE), a is the coefficient of converting electromotive force into metabolic energy, V is the volume of the plant cells, P is the pressure to which the plant cells are subjected, and the pressure P is expressed by a pressure formulaCalculating 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 intoDue 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 capacitance of the plant leaf changing along with the clamping force can be deformed intoWherein 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 equationDeduced, wherein Xl is a physiological inductive reactance, E is an electromotive force, E0Is a standard electromotive force, R0Is the ideal gas constant, T is the temperature, MiDielectric concentration, M, in response to physiological inductance within the cell membraneoConcentration of dielectric substances, L, in response to physiological inductance outside the cell membrane0Dielectric substance concentration M being responsive to physiological inductive reactance in cell membraneiThe 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 reactanceT=Mi+Mo,F0Is the Faraday constant, nXLIs the number of dielectric mass transfers in response to physiological inductance; e can be used for doing work, PV is proportional to PV (a is aE), a is the coefficient of converting electromotive force into metabolic energy, V is the volume of the plant cells, P is the pressure to which the plant cells are subjected, and the pressure P is expressed by a pressure formulaCalculating 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 intoDue 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 intoWherein q is0、k3And b3Are parameters of the model.
Further, in the step ten, the dielectric substance transfer number Kn of the k-type response physiological resistance is obtained according to the parameters in the model in the step sevenRThe method comprises the following steps: knR=lnk1-lny0B type dielectric transition number Bn in response to physiological resistanceRThe method comprises the following steps: bnR=b1。
Further, in the eleventh step, the dielectric substance transfer number Kn of the k-type response physiological capacitive reactance is obtained according to the parameters in the model in the eighth stepXCThe method comprises the following steps: knXC=lnk2-lnp0B type dielectric transition number Bn in response to physiological capacitive impedanceXCThe method comprises the following steps: bnXC=b2。
Further, in the twelfth step, the dielectric material transfer number Kn of the k-type response physiological inductive reactance is obtained according to the parameters in the ninth stepXLThe method comprises the following steps: knXL=lnk3-lnq0B type dielectric transition number Bn in response to physiological inductive reactanceXLThe method comprises the following steps: bnXL=b3。
Further, the k-type total dielectric material transfer number Kn in the third stepTThe acquisition method comprises the following steps: knT=KnR+KnXC+KnXL。
Further, in the fourteenth stepb type total dielectric transition number BnTThe acquisition method comprises the following steps: bnT=BnR+BnXC+BnXL。
Further, the percentage of dielectric mass transfer KPn for k-type response to physiologic resistance in step fifteenRThe calculation method comprises the following steps:unit%; percentage of dielectric mass transfer KPn for k-type response to physiological capacitive impedanceXCThe calculation method comprises the following steps:unit%; percent dielectric transfer KPn of k-type response to physiological susceptibilitiesXLThe calculation method comprises the following steps:the unit% is.
Further, the type b dielectric substance transfer percentage BPn responding to physiological resistance in the step sixteenRThe calculation method comprises the following steps:unit%; b-type dielectric substance transfer percentage BPn responding to physiological capacitive reactanceXCThe calculation method comprises the following steps:unit%; b-type dielectric substance transfer percentage BPn responding to physiological inductive reactanceXLThe calculation method comprises the following steps:the unit% is.
The invention has the following beneficial effects:
1. the method can rapidly and quantitatively detect the transfer percentages of various types of dielectric substances of different plant leaves under different environments on line, determine the contribution parts of phospholipids, surface proteins (peripheral proteins) and binding proteins (intrinsic proteins) on cell membranes to the operation of cell membrane substances, and the determination results are comparable.
2. The invention characterizes the water and substance transport capacity of leaf cells and the communication characteristics of different plant leaf water and substances in a system under different environments by measuring the transfer condition of dielectric substances of the plant leaf cells.
3. The invention is simple and convenient, has wide applicability and low price of required instruments.
Drawings
FIG. 1 is a structural model of cell membrane
FIG. 2 is a schematic structural view 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 capacitance of plant leaf, C is the physiological capacitance of plant leaf, 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 Xl is the physiological inductive reactance of the plant leaf, Xc is the physiological capacitive reactance of the plant leaf, Z is the physiological impedance of the plant leaf, and R is the physiological resistance of the plant leaf.
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 a Nernst equation, the physiological resistance is inversely proportional to the conductivity, and the conductivity is directly 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 (1):
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, in K; ciConcentration of dielectric substances, C, in response to physiological resistance in cell membranesoThe 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 membraneT=Ci+Co,F0Is the Faraday constant, equal to 96485C.mol-1;nRIs the number of dielectric material transitions 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, CoAnd CiThe sum is constant and equal to the total amount C of dielectric substances responding to physiological resistance inside and outside the membraneT,CiIt is proportional to the conductivity, which is the inverse of the resistance R, and, therefore,can be expressed asWherein R is resistance, f0Concentration C of dielectric substance responsive to physiological resistance in cell membraneiAnd the proportionality coefficient of the conversion between resistance, therefore, (2) can become:
(3) is transformed to obtain
(4) Is transformed to obtain
(5) 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 (7) is a physiological resistance due to the specific effective thickness of the plant leaves(7) The formula can be deformed into:
d, a and E in formula (7) for the same blade to be tested in the same environment0、R0、T、nR、F0、CT、f0Are all constant values; order toThus, equation (8) can be transformed into:
(9) in the formula y0、k1And b1Are parameters of the model.In this case, the amount of the solvent to be used,e in (A)0、R0T and F0Is a constant value, and the value is,dielectric substance transfer responsive only to physiological resistanceNumber nRIn direct proportion to the total weight of the steel, and in the same way,in this case, the amount of the solvent to be used,d, a, R in (1)0T and F0Is a constant value, and the value is,the same holds only for the number of dielectric material transitions n in response to physiological resistanceRProportional, therefore, k-type dielectric material transfer number Kn in response to physiological resistanceR=lnk1-lny0B type dielectric transition number Bn in response to physiological resistanceR=b1。
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 Nernst 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 follows (10):
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; qiDielectric substance concentration, Q, in response to physiological capacitive reactance in cellular membranesoThe total amount of dielectric substance Q ═ Q for responding to physiological capacitive reactance outside cell membrane and for responding to physiological capacitive reactance outside membranei+Qo,F0Is the Faraday constant, equal to 96485C.mol-1;nXCIs 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, QoAnd QiThe sum is certain and equal to the total amount of dielectric substances Q, Q responding to physiological capacitive reactance inside and outside the membraneiIs 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 asXc is capacitive reactance, J0Is a physiological capacitive reactance in cell membraneDielectric substance concentration Q ofiAnd capacitive reactance, and thus (11) may become:
(12) is transformed to obtain
(13) Can become:
(14) 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 (16) is a physiological capacitive reactance due to the specific effective thickness of the plant leaves(16) The formula can be deformed into:
for the same blade to be measured in the same environment, (17) formula (d, a, E)0、R0、T、nXC、F0、Q、J0Are all constant values, orderTherefore, equation (17) can be transformed into:
(18) in the formula p0、k2And b2Are parameters of the model.In this case, the amount of the solvent to be used,e in (A)0、R0T and F0Is a constant value, and the value is,number n of dielectric substance transitions responsive only to physiological capacitive impedanceXCIn direct proportion to the total weight of the steel, and in the same way,in this case, the amount of the solvent to be used,d, a, R in (1)0T and F0Is a constant value, and the value is,number n of dielectric substance transitions which likewise only respond to physiological capacitive impedanceXCProportional, k-type dielectric material transfer number Kn in response to physiological capacitive impedanceXC=lnk2-lnp0B type dielectric transition number Bn in response to physiological capacitive impedanceXC=b2。
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 (19):
wherein E is electromotive force, E0Is a standard electromotive force, R0Is an ideal gasNumber equal to 8.314570J.K-1.mol-1(ii) a T is temperature, in K; miDielectric concentration, M, in response to physiological inductance within the cell membraneoThe total amount of dielectric substance M is the concentration of dielectric substance responding to physiological inductance outside the cell membraneT=Mi+Mo,F0Is the Faraday constant, equal to 96485C.mol-1;nXLIs 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, MoAnd MiThe 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 membraneT,MiIt is proportional to the conductivity of the dielectric material in response to the physiological impedance, which is the inverse of the impedance Xl, and therefore,can be expressed asXl is inductive reactance, L0Dielectric substance concentration M being responsive to physiological inductive reactance in cell membraneiAnd inductance, and thus, the (20) formula may become:
(21) is transformed to obtain
(22) Can become:
(23) the two sides of the formula are taken as indexes and can be changed into:
further modified, it is possible to obtain:
in the formula (25), Xl is a physiological inductive reactance due to the specific effective thickness of the plant leaves(25) The formula can be deformed into:
for the same blade to be measured under the same environment, (26) formula (d, a, E)0、R0、T、nXL、F0、MT、L0Are all constant values, orderThus, equation (26) can be transformed as:
(27) in the formula q0、k3And b3Are parameters of the model.In this case, the amount of the solvent to be used,e in (A)0、R0T and F0Is a constant value, and the value is,number n of dielectric material transitions responsive only to physiological inductive reactanceXLIn direct proportion to the total weight of the steel, and in the same way,in this case, the amount of the solvent to be used,d, a, R in (1)0T and F0Is a constant value, and the value is,number n of dielectric material transitions which likewise only respond to physiological inductive reactanceXLProportional, therefore, k-type dielectric material transfer number Kn in response to physiological inductive reactanceXL=lnk3-lnq0B type dielectric transition number Bn in response to physiological inductive reactanceXL=b3。
A measuring device for measuring the transfer number of dielectric substances of plant leaf cells is shown in figure 2 and 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; 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 (HIOKI3532-50 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.
Examples
For example, bletilla striata. Picking biennial rhizoma bletillae plants in a base of a Prading karst ecological comprehensive test station in Guizhou province of Chinese academy of sciences, quickly returning to a laboratory, cleaning surface dust of leaves on fresh branches, respectively collecting the leaves to be tested from the fresh branches one by one, and soaking the leaves in distilled water for 30 minutes; sucking water on the surface of the leaf, immediately clamping the leaf to be measured between parallel electrode plates of a measuring device, setting measuring voltage and frequency, setting different clamping forces by changing the mass of an iron block, and measuring physiological capacitance, physiological resistance and physiological impedance of the plant leaf under different clamping forces in a parallel mode; the physiological capacitance of different blades and different clamping forces are shown in table 1, the physiological resistance is shown in table 2, and the physiological impedance is shown in table 3. Calculating physiological capacitive reactance according to the data in table 1 as shown in table 4, and calculating physiological inductive reactance of plant leaves according to the data in table 2, table 3 and table 4 as shown in table 5; a model of the physiological resistance of the plant leaf changing with the clamping force is constructed according to the data in the table 2, such as the table 6, and a model of the physiological capacitive reactance of the plant leaf changing with the clamping force is constructed according to the data in the table 4, such as the table 7. A model of the physiological sensory resistance of the plant leaf as a function of the clamping force was constructed from the data in Table 5 as shown in Table 8. According to the parameters of each model in Table 6, the dielectric material transfer number Kn of k-type response physiological resistance is obtainedRAnd b-type dielectric material transfer number Bn responding to physiological resistanceR(see Table 9); according to the parameters of the models in Table 7, respectivelyDielectric substance transfer number Kn for obtaining k-type response physiological capacitive impedanceXCAnd b-type dielectric substance transfer number Bn in response to physiological capacitive impedanceXC(Table 9); according to the parameters of each model in Table 8, the dielectric material transfer number Kn of k-type response physiological inductive reactance is obtainedXLAnd b-type dielectric material transfer number Bn responding to physiological inductive reactanceXL(Table 9); according to KnR、KnXCAnd KnXLObtaining the k-type total dielectric material transfer number KnT(Table 9); according to BnR、BnXCAnd BnXLObtaining the b-type total dielectric material transfer number BnT(Table 9); according to KnR、KnXC、KnXLAnd KnTObtaining KPn dielectric mass transfer percentages of k-type response physiological resistanceRDielectric mass transfer percentage of k-type response to physiological capacitive reactance KPnXCAnd percent dielectric transfer of k-type responsive physiological susceptibilities KPnXL(Table 10); according to BnR、BnXC、BnXLAnd BnTObtaining the dielectric substance transfer percentage BPn of the b-type response physiological resistance respectivelyRType b dielectric mass transfer percentage BPn in response to physiological capacitive impedanceXCAnd b-type dielectric mass transfer percentage BPn in response to physiological inductanceXL(Table 10).
TABLE 1 physiological capacitance (pF) of different rhizoma Bletillae leaf positions under different clamping force (F, unit N)
TABLE 2 physiological resistance (M omega) of different bletilla striata leaf at different leaf positions under different clamping force (F, unit N)
TABLE 3 physiological impedance (M Ω) of different leaf positions and rhizoma Bletillae under different clamping force (F, unit N)
TABLE 4 physiological capacitive reactance (M omega) of different rhizoma Bletillae leaf positions under different clamping forces (F unit N)
TABLE 5 physiological inductive reactance (M omega) of different rhizoma Bletillae leaf positions under different clamping forces (F, unit N)
TABLE 6 model (R-F) of physiological resistance (R) of different leaf positions and leaves with clamping force (F) and parameters
TABLE 7 physiological capacitive reactance (Xc) of different leaf positions and leaves of bletilla striata model (Xc-F) and parameters thereof
TABLE 8 physiological inductance (Xl) of different leaf positions and leaves with the change model (Xl-F) of clamping force (F) and parameters
TABLE 9 dielectric transition number Kn of k-type response to physiological resistance of leaves at different leaf positionsRDielectric substance transfer number Kn in response to physiological capacitive impedanceXCDielectric substance transfer number Kn in response to physiological inductanceXLK type total dielectric material transfer number KnTAnd b-type dielectric material transfer number Bn responding to physiological resistanceRDielectric substance transfer number Bn in response to physiological capacitive impedanceXCDielectric material transfer number Bn in response to physiological inductanceXLB type total dielectric material transfer number BnT
TABLE 10 percent dielectric transfer KPn of k-type response to physiologic resistance for different leaf positionsR(%), percent dielectric transfer in response to physiological capacitive impedance KPnXC(%), percent dielectric transfer in response to physiological inductance KPnXL(%) and b-type dielectric transfer percentage in response to physiological resistance BPnR(%), percent dielectric transfer in response to physiological capacitive impedance, BPnXC(%) and percent dielectric transfer in response to physiological inductance BPnXL(%)
The implementation effect of the invention is as follows:
as can be seen from table 10, the percentage of dielectric substance transfer of the same type of bletilla striata leaves at different positions is different regardless of the k-type or b-type, but the percentage of dielectric substance transfer of the same type of bletilla striata leaves at the same position is smaller. In addition, it can be seen from Table 10 that the percent transfer of dielectric species in response to physiological resistance is less different from the percent transfer of dielectric species in response to physiological impedance, and in general, the percent transfer of dielectric species in response to physiological resistance is greater than the percent transfer of dielectric species in response to physiological impedance, which may be related to the fact that the percent transfer of dielectric species in response to physiological resistance reflects passive transport, and the percent transfer of dielectric species in response to physiological impedance reflects carrier transport, with the two species flowing in opposite directions, with the charge approaching equilibrium, and with the weakly negative charge remaining outside the membrane.
The percentage of dielectric substance transfer in response to physiological capacitive reactance is related to the potential difference between the inside and the outside of the membrane, while the potential difference between the inside and the outside of the membrane is related to the concentration difference of substances inside and outside the membrane, and the larger the potential difference between the inside and the outside of the membrane, the larger the concentration of substances inside the membrane is, the more substances can be transferred; thus, with respect to the source library relationship, the greater the percent dielectric mass transfer in response to physiological capacitive reactance, the greater the effect as a source and vice versa. As can be seen from table 10, the leaf source in the fifth leaf position had a greater effect than the fourth and third leaf positions, and the leaf bank in the first and second leaf positions had a greater effect than the third and fourth leaf positions with respect to moisture and stored organic nutrients. This indicates that the young leaves need to obtain moisture and nutrients from the mature leaves, which have the characteristics of the sink compared to the mature leaves, which have the characteristics of the source. Compared with the single-head bletilla striata tissue culture seed stem, the saddle bletilla striata tissue culture seed stem accelerates the growth and development of new bletilla striata leaves due to twice source-to-reservoir ratio, so that the production has more advantages. In addition, high yielding bletilla striata all have the characteristics of enlarged basal leaves and late abscission, which can be explained by the examples of the present invention.
Although the embodiments of the present invention have been described with reference to the accompanying drawings, it is not intended to limit the scope of the present invention, and it should be understood by those skilled in the art that various modifications and variations can be made without inventive efforts by those skilled in the art based on the technical solution of the present invention.
Claims (7)
1. A method for determining the dielectric substance transfer number of plant leaf cells is characterized by comprising the following steps:
step one, connecting a measuring device with an LCR tester;
selecting a fresh branch of a plant to be detected, wrapping the base of the branch, and bringing the branch to a laboratory;
collecting leaves to be detected from the fresh branches, 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, respectively obtaining the dielectric substance transfer number Kn of the k-type response physiological resistance according to the parameters in the model in the step sevenRAnd b-type dielectric material transfer number Bn responding to physiological resistanceR;
Step eleven, respectively obtaining the dielectric substance transfer number Kn of the k-type response physiological capacitive reactance according to the parameters in the model in the step eightXCAnd b-type dielectric substance transfer number Bn in response to physiological capacitive impedanceXC,KnXC=lnk2-lnp0B type dielectric transition number Bn in response to physiological capacitive impedanceXCThe method comprises the following steps: bnXC=b2;
Step twelve, respectively obtaining the dielectric substance transfer number Kn of the k-type response physiological inductive reactance according to the parameters in the model in the step nineXLAnd b-type dielectric material transfer number Bn responding to physiological inductive reactanceXL,KnXL=lnk3-lnq0B type dielectric transition number Bn in response to physiological inductive reactanceXLThe method comprises the following steps: bnXL=b3;
Thirteen, responding to the dielectric substance transfer number Kn of the physiological resistance according to the k typeRK-type dielectric substance transfer number Kn of response physiological capacitive reactanceXCAnd k-type dielectric substance transfer number Kn in response to physiological inductive reactanceXLObtaining the k-type total dielectric material transfer number KnT,KnT=KnR+KnXC+KnXL;
Fourteen, responding to the dielectric material transfer number Bn of the physiological resistance according to type bRType b dielectric material transfer number Bn responding to physiological capacitive impedanceXCAnd b-type dielectric material transfer number Bn responding to physiological inductive reactanceXLObtaining the b-type total dielectric material transfer number BnT,BnT=BnR+BnXC+BnXL;
Step fifteen, according to KnR、KnXC、KnXLAnd KnTObtaining KPn dielectric mass transfer percentages of k-type response physiological resistanceRDielectric mass transfer percentage of k-type response to physiological capacitive reactance KPnXCAnd percent dielectric transfer of k-type responsive physiological susceptibilities KPnXL,Unit%;unit%; unit%;
step sixteen, according to BnR、BnXC、BnXLAnd BnTObtaining the dielectric substance transfer percentage BPn of the b-type response physiological resistance respectivelyRType b dielectric mass transfer percentage BPn in response to physiological capacitive impedanceXCAnd b-type dielectric mass transfer percentage BPn in response to physiological inductanceXL,Unit%;unit%;unit%; in the seventh step, the physiological resistance of the plant leaf changes along with the clamping force,the model is based on the Nernst equationDerived, wherein R is resistance, E is electromotive force, E is0Is a standard electromotive force, R0Is an ideal gas constant, T is temperature, CiConcentration of dielectric substances, C, in response to physiological resistance in cell membranesoConcentration of dielectric substances in response to physiological resistance outside cell membrane, f0Concentration C of dielectric substance responsive to physiological resistance in cell membraneiProportional coefficient converted from physiological resistance, total dielectric substance C of intra-membrane and extra-membrane response physiological resistanceT=Ci+Co,F0Is the Faraday constant, nRIs the number of dielectric mass transfers in response to physiological resistance; e is proportional to PV, PV is a E, a is the coefficient of conversion of electromotive force to metabolic energy, V is plant cellVolume, P is the pressure to which the plant cells are subjected, and pressure P is expressed by the pressure equationCalculating 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 intoDue 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 resistance of the plant leaf changing along with the clamping force can be deformed intoWherein y is0、k1And b1Is a parameter of the model; in the eighth step, the physiological capacitive reactance of the plant leaves is clampedThe model of the change of the holding force is,the model is based on the Nernst equation Deduced, wherein Xc is capacitive reactance, E is electromotive force, E0Is a standard electromotive force, R0Is the ideal gas constant, T is the temperature, QiDielectric substance concentration, Q, in response to physiological capacitive reactance in cellular membranesoConcentration 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 membranesiThe 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 Qi+Qo,F0Is the Faraday constant, nXCIs the number of dielectric mass transfers in response to physiological capacitive reactance; e is proportional to PV, and PV is a E, a is the coefficient of conversion of electromotive force to metabolic energy, V is the volume of plant cell, P is the pressure to which plant cell is exposed, and the pressure P is expressed by the pressure equationCalculating 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 intoDue to the specific effective thickness of the plant leavesTherefore, 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 intoWherein p is0、k2And b2Is a parameter of the model; in the ninth step, the physiological inductive reactance of the plant leaf changes along with the clamping force, the model is based on the Nernst equationDeduced, wherein Xl is inductive reactance, E is electromotive force, E0Is a standard electromotive force, R0Is the ideal gas constant, T is the temperature, MiDielectric concentration, M, in response to physiological inductance within the cell membraneoConcentration of dielectric substances, L, in response to physiological inductance outside the cell membrane0Dielectric substance concentration M being responsive to physiological inductive reactance in cell membraneiProportionality coefficient of conversion between physiological inductive reactanceTotal amount of dielectric substance M for intra-and extramembranous response to physiological inductive reactanceT=Mi+Mo,F0Is the Faraday constant, nXLIs the number of dielectric mass transfers in response to physiological inductance; e is proportional to PV, and PV is a E, a is the coefficient of conversion of electromotive force to metabolic energy, V is the volume of plant cell, P is the pressure to which plant cell is exposed, and the pressure P is expressed by the pressure equationCalculating 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 intoDue 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 intoWherein q is0、k3And b3Are parameters of the model.
2. The method for determining the number of dielectric substance transfers in plant leaf cells according to claim 1, wherein: survey device includes support (1), cystosepiment (2), plate electrode (3), wire (4), iron plate (5), plastic rod (6) and fixation clamp (7), support (1) is the rectangular frame structure, and one side is open, open on support (1) upper end has the through-hole, supply plastic rod (6) to stretch into, support (1) lower extreme inwards one side and plastic rod (6) bottom are stained with two cystosepiments (2) respectively, inlay plate electrode (3) in cystosepiment (2), a wire (4) are drawn forth respectively in two plate electrodes (3), can place iron plate (5) of fixed quality on cystosepiment (2) of plastic rod (6), plastic rod (6) are located the inside one end of support and are fixed by fixation clamp (7).
3. The method for determining the number of dielectric substance transfers in plant leaf cells according to claim 2, wherein: the electrode plate (3) is a circular electrode plate, and the electrode plate (3) is made of copper.
4. The method for determining the number of dielectric substance transfers in plant leaf cells according to claim 2, wherein: 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.
5. The method for determining the number of dielectric substance transfers in plant leaf cells according to claim 1, wherein: in the fifth step, a calculation formula of the physiological capacitive reactance of the plant leaves is as follows:wherein Xc is the physiological capacitance of plant leaf, C is the physiological capacitance of plant leaf, f is the test frequency, and pi is the circumference ratio equal to3.1416。
6. The method for determining the number of dielectric substance transfers in plant leaf cells according to claim 1, wherein: in the sixth step, a calculation formula of the physiological inductive reactance of the plant leaf is as follows:wherein Xl is the physiological inductive reactance of the plant leaf, Xc is the physiological capacitive reactance of the plant leaf, Z is the physiological impedance of the plant leaf, and R is the physiological resistance of the plant leaf.
7. The method for determining the number of dielectric substance transfers in plant leaf cells according to claim 1, wherein: in the step ten, the dielectric substance transfer number Kn of the k-type response physiological resistance is obtained according to the parameters in the model in the step sevenRThe method comprises the following steps: knR=lnk1-lny0B type dielectric transition number Bn in response to physiological resistanceRThe method comprises the following steps: bnR=b1。
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