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Research article Special Issues

Characterization of Lysobacter enzymogenes B25, a potential biological control agent of plant-parasitic nematodes, and its mode of action

  • Received: 09 December 2022 Revised: 16 February 2023 Accepted: 22 February 2023 Published: 01 March 2023
  • It is certainly difficult to estimate productivity losses due to the action of phytopathogenic nematodes but it might be about 12 % of world agricultural production. Although there are numerous tools to reduce the effect of these nematodes, there is growing concern about their environmental impact. Lysobacter enzymogenes B25 is an effective biological control agent against plant-parasitic nematodes, showing control over root-knot nematodes (RKN) such as Meloidogyne incognita and Meloidogyne javanica. In this paper, the efficacy of B25 to control RKN infestation in tomato plants (Solanum lycopersicum cv. Durinta) is described. The bacterium was applied 4 times at an average of concentration around 108 CFU/mL showing an efficacy of 50–95 % depending on the population and the pressure of the pathogen. Furthermore, the control activity of B25 was comparable to that of the reference chemical used. L. enzymogenes B25 is hereby characterized, and its mode of action studied, focusing on different mechanisms that include motility, the production of lytic enzymes and secondary metabolites and the induction of plant defenses. The presence of M. incognita increased the twitching motility of B25. In addition, cell-free supernatants obtained after growing B25, in both poor and rich media, showed efficacy in inhibiting RKN egg hatching in vitro. This nematicidal activity was sensitive to high temperatures, suggesting that it is mainly due to extracellular lytic enzymes. The secondary metabolites heat-stable antifungal factor and alteramide A/B were identified in the culture filtrate and their contribution to the nematicidal activity of B25 is discussed. This study points out L. enzymogenes B25 as a promising biocontrol microorganism against nematode infestation of plants and a good candidate to develop a sustainable nematicidal product.

    Citation: Sònia Martínez-Servat, Lola Pinyol-Escala, Oriol Daura-Pich, Marta Almazán, Iker Hernández, Belén López-García, Carolina Fernández. Characterization of Lysobacter enzymogenes B25, a potential biological control agent of plant-parasitic nematodes, and its mode of action[J]. AIMS Microbiology, 2023, 9(1): 151-176. doi: 10.3934/microbiol.2023010

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  • It is certainly difficult to estimate productivity losses due to the action of phytopathogenic nematodes but it might be about 12 % of world agricultural production. Although there are numerous tools to reduce the effect of these nematodes, there is growing concern about their environmental impact. Lysobacter enzymogenes B25 is an effective biological control agent against plant-parasitic nematodes, showing control over root-knot nematodes (RKN) such as Meloidogyne incognita and Meloidogyne javanica. In this paper, the efficacy of B25 to control RKN infestation in tomato plants (Solanum lycopersicum cv. Durinta) is described. The bacterium was applied 4 times at an average of concentration around 108 CFU/mL showing an efficacy of 50–95 % depending on the population and the pressure of the pathogen. Furthermore, the control activity of B25 was comparable to that of the reference chemical used. L. enzymogenes B25 is hereby characterized, and its mode of action studied, focusing on different mechanisms that include motility, the production of lytic enzymes and secondary metabolites and the induction of plant defenses. The presence of M. incognita increased the twitching motility of B25. In addition, cell-free supernatants obtained after growing B25, in both poor and rich media, showed efficacy in inhibiting RKN egg hatching in vitro. This nematicidal activity was sensitive to high temperatures, suggesting that it is mainly due to extracellular lytic enzymes. The secondary metabolites heat-stable antifungal factor and alteramide A/B were identified in the culture filtrate and their contribution to the nematicidal activity of B25 is discussed. This study points out L. enzymogenes B25 as a promising biocontrol microorganism against nematode infestation of plants and a good candidate to develop a sustainable nematicidal product.



    Differential line geometry examines the set of lines in space with 3-dimension. The ambient space can be an Euclidean or a non-Euclidean space. It is one of the interesting subdivision of differential geometry since it is directly connected with the spatial motion (kinematics) and get therefore a lot of application in mechanism design [1,2,3,4,5,6,7,8,9].

    A line congurence is a two-parameter set of lines in three-dimensional space- is a classical part of differential geometry, whose origins can be traced to Kummer in his memoir; Allgemeine Theorie der Gradlinigen Strahen system [1]. The normal vector field of a surface could be given as an example of line congruence. In this way the line congruence formed by the normal vector field of the surface constructs a special class, which is called normal line congruence. However, the line congruence does not usually need to consist only of normal vector field. The lines of the line congruence which pass through a curve on the surface form a one-parameter family of lines in the space or ruled surface (parameter ruled surface). Recently, line congruence has become relevant for practical applications (See for instance [2,3,4,5,6,7]).

    As it is known, the most analytical tool in the study of three-dimensional kinematics and differential line geometry is based upon the so-called E. Study's map: The set of all oriented lines in Euclidean 3-space E3 is in one-to-one correspondence with set of points of the dual unit sphere in the dual 3-space D3. It allows a complete generalization for the spherical point geometry to the spatial line geometry by means of dual-number extension, i.e. replacing all ordinary quantities by the corresponding dual-number quantities. Hence, the ruled surfaces and line congruence obtained by the motion of a line, depending on one and two parameters, respectively. There exists a vast literature on the subject including several monographs, such as [6,8,9,10,11].

    This work develops the kinematic-geometry for line congruence in the space of lines by using the analogy with theory of surfaces. Then the well known formulae of J. Liouville, Hamilton, and Mannheim of surfaces theory are proved for the line congruence. Moreover, a new geometrical interpretation of J. Liouville formula has been defined for a given closed ruled surface in the line congruence. The Plücker conoid associated with the line congruence has been derived and it is shown that the principal axes of it are located at its center and at right angles. Finally, an example of application is investigated and explained in detail.

    In this section we list some notions, formulas of dual numbers and dual vectors (See for instance [1,2,3,4,5,6]). Therefore we start with recalling the use of appropriate line coordinates: An oriented line L in the Euclidean 3-space E3 can be determined by a point pL and a normalized direction vector x of L, i.e. x=1. To obtain components for L, one forms the moment vector

    x=p×x (2.1)

    with respect to the origin point in E3. If p is substituted by any point

    q=p+μx,μR (2.2)

    on L, then Eq (2.1) implies that x is independent of p on L. The two vectors x and x are not independent of one another; they satisfy the following relationships:

    <x,x>=1,<x,x>=0. (2.3)

    The six components xi,xi(i=1,2,3) of x and xare called the normalized Plűcker coordinates of the line L. Hence the two vectors x and x determine the oriented line L.

    In line geometry, from a kinematic point of view, a line congruence (congruence for shortness) is a two-parameter set of lines in E3 generated by a straight line L moving along a surface. The various positions of the generating lines are called the rulings or generators of the congruence. This congruence holds a parameterization as form

    Q:Y(u1,u2,v)=y(u1,u2)+vr(u1,u2),vR. (2.4)

    Here y=y(u1,u2) is called the director surface, r=r(u1,u2) is the unit vector giving the direction of generating line of the congruence, u1, u2 are the motion parameters, and v is the parameter of its points indicating the singed distance of the corresponding point on y=y(u1,u2). The equations

    u1=u1(t),u2=u2(t),u21+u220,tR, (2.5)

    define a ruled surface belonging to the congruence. It is a developable if and only if the determinate

    (r,dy,dr)=0. (2.6)

    Ruled surfaces (such as cylinders and cones) contain rulings where the tangent plane touches the surface along the entire line. Such rulings are called torsal lines, to distinguish them from the common case of the non-torsal generators [6,11,12,13]:

    Ruled surfaces with exclusively torsal generators are called developable surfaces;

    Ruled surfaces consisting largely of non-torsal generators are called skew ruled surfaces (or warped ruled surfaces);

    Cylinders, cones, and ruled surfaces that consist of tangents of a spatial curve are developable surfaces.

    Definition 1. The singular surface points of a torsal generator is called its cuspidal point, and the tangent plane in its other direction is called torsal plane.

    The set of dual numbers is

    D={X=x+εx x, xR, ε0, ε2=0}. (2.7)

    This set is a commutative ring under addition and multiplication. This set cannot be a field under these operations, because 0+εx has no multiplication inverse in D. But this ring has a unit element according to multiplication. A dual number X=x+εx, is called proper if x0. An example of dual number is the dual angle subtended by two skew lines in the Euclidean 3-space E3 and defined as Θ=ϑ+εϑ in which ϑ and ϑ are, respectively, the projected angle and the minimal distance between the two lines.

    For all pairs (x,x)E3×E3 the set

    D3={X=x+εx,ε0,ε2=0}, (2.8)

    together with the scalar product

    <X,Y>=<x,y>+ε(<y,x>+<y,x>), (2.9)

    forms the dual 3-space D3.Thereby a point X=(X1,X2,X3)t has dual coordinates Xi=(xi+εxi)D. The norm is defined by

    <X,X>12:=X=x(1+ε<x,x>x2). (2.10)

    In the dual 3-space D3 the dual unit sphere is defined by

    K={XD3X2=X21+X22+X23=1}. (2.11)

    Definition of dual unit sphere gives us that all points X of K must satisfy two equations

    x21+x22+x23=1,x1x1+x2x2+x3x3=0. (2.12)

    Via this we have the following map (E. Study's map): The set of all oriented lines in the Euclidean 3-space E3 is in one-to-one correspondence with the set of points of dual unit sphere in the dual 3-space D3. By using this correspondence, one can derive the properties of the spatial motion of a line. Hence, the geometry of ruled surface is represented by the geometry of curves on the dual unit sphere in D3. Therefore, the dual curve and ruled surfaces are synonymous in this paper.

    The E. Study's map admits us to revision Eq (2.4) using the dual vector function as

    R(u1,u2)=r(u1,u2)+εy(u1,u2)×r(u1,u2)=r(u1,u2)+εr(u1,u2), (2.13)

    where r is the moment of r about the origin in E3. Since the spherical image r(u1,u2) is a unit vector, then the dual vector R(u1,u2) also has unit magnitude as is seen from the computations:

    <R,R>=<r,r>+2ε<r,y×r>+ε2<y×r,y×r>=<r,r>=1.

    Thus the line congruence fills a domain on dual unit sphere K in D3. Hence, the line congruence can be viewed as a two-dimensional surface in D3-space. It follows that there are resemblances between theory of surface and theory of line congruence.

    A relationship such as f(u1,u2)=0 between the parameters u1, u2 reduces the congruence to a one-parameter set of lines (a ruled surface) in the congruence R(u1,u2). The equations u2=c2 and  u1=c1(real constants) determines parameter ruled surfaces given by the parameter dual curves R=R(u1,c2) and R=R(c1,u2), respectively.

    Definition 2. A line congruence is torsal, if the ruled surfaces defined by u1=const. are developable, and so are the ones defined by u2=const.

    If L any function, scalar or vector, defined for the line congruence, we shall denote Lu1, and Lu2 by L/u1, and L/u2, respectively. Thus Ru1 is a dual tangent vector in direction which u1 alone varies, similarly for Ru2. Therefore,

    <Ru1,Ru1>=E=e+εe,<Ru1,Ru2>=F=f+εf,<Ru2,Ru2>=G=g+εg.} (2.14)

    Thus, we arrive by means of the real and dual parts of Eq (2.14), at

    e=<ru1,ru1>,e=2<ru1,ru1>,f=<ru1,ru2>,f=<ru1,ru2>+<ru1,ru2>,g=<ru2,ru2>,g=2<ru2,ru2>.} (2.15)

    Now we consider two neighboring dual points, with position vectors R and R+dR, from (u1,u2) to (u1+du1,u2+du2), respectively. Then

    dR=Ru1du1+Ru2du2. (2.16)

    Since the two dual points are adjacent points on a dual curve passing through them, the dual arc length dS(=ds+εds) is

    dS2=<dR,dR>=<Ru1du1+Ru2du2,Ru1du1+Ru2du2>,

    or finally,

    dS2=Edu21+2Fdu1du2+Gdu22. (2.17)

    By separating the real and dual parts of this equation, respectively, we get

    I(u1,u2):=ds2=<dr,dr>=edu21+2fdu1du2+gdu22, (2.18)

    and

    II(u1,u2):=2dsds=<dr,dr>=edu21+2fdu1du2+gdu22. (2.19)

    The forms I and II are called the first and second fundamental forms of the line congruence, respectively. The distribution parameter of a ruled surface belong to the congruence is given by [10,11]

    λ=12(III)=edu21+2fdu1du2+gdu22edu21+2fdu1du2+gdu22. (2.20)

    A ruled surface belong to the congruence for which

    edu21+2fdu1du2+gdu22=0 (2.21)

    is a developable surface. If we let η=du1/du2, we can write

    eη2+2fη+g=0. (2.22)

    For developable surfaces of the congruence, we equate the coefficients of the Eq (2.22) to zero identically, and we can write

    e=2<ru1,ru1>=0,f=<ru1,ru2>+<ru1,ru2>=0,g=2<ru2,ru2>=0.} (2.23)

    Through every line in the congruence there pass two principal surfaces whose images on the dual unit sphere we take as parameter curves. For this special system we assume that u1, and u2 dual curves of R(u1,u2) are principal ruled surfaces, i.e., the elements f and f of the first and second fundamental forms vanish identically (f=f=0). So, according to Eq (2.17), the dual arc length of the dual curves u2=c2(real const.), and u1=c1(real const.), respectively are dS1=ds1+εds1=Edu1 and dS2=ds2+εds2=Gdu2.

    In order to examine the geometrical properties of the congruence, we set up a moving frame coincident with the point on the sphere. If we label the point on the sphere, the generator as R(u1,u2), then the Blaschke frame can be set up [10,11,15]

    R12=Ru1Ru1=Ru1E, R22=Ru2Ru2=Ru1G, R=R12×R22, (2.24)

    which are invariants vector functions on the congruence Q; we fix det(R, R12,R22)=+1 and consequently

    λ=12(edu21+gdu22edu21+gdu22)=e2(du1ds)2+g2(du2ds)2. (2.25)

    The extreme values of the distribution parameter, corresponding to the principal surfaces R(u1,c2), and R(c1,u2), respectively, are obtained by

    λ1:=ds1ds1=e2e, λ2:=ds2ds2=g2g. (2.26)

    According to the elements of spherical kinematics, the motion of the frame {O; R, R12, R22} at any instant is a rotation around the Darboux vector of this frame. Hence, by means of the derivatives with respect to the dual arc-length parameter of the dual curves u2=c2 with tangent R12, the derivative formula is [10,15]

    S1(RR12R22)=(  0  101  0Σ1  0Σ10)(RR12R22)=Ω1×(RR12R22), (2.27)

    where

    Ω1=Σ1R+R22. (2.28)

    Σ1=σ1+εσ1=Eu22EG is the geodesic curvature of the dual curves R(u1,c2). Similarly, the derivative formula of the Blaschke frame of the dual curves R(c1,u2), with tangent R22 is

    S2(RR12R22)=(  0  01  0  0Σ21Σ20)(RR12R22)=Ω2×(RR12R22), (2.29)

    where

    Ω2=Σ2RR12. (2.30)

    Σ2=σ2+εσ2=Gu12GE has the same meaning as in Eq (2.27). In other words, dual geodesic curvatures of the dual curves R(u1,c2), and R(c1,u2), respectively, are

    Σi=σi+εσi=det(R,dRdSi,d2RdS2i),(i=1,2). (2.31)

    Consider ui=ui(t), (i=1,2) as functions of real parameter tR. Then R=R(u1(t),u2(t))represents a ruled surface in the congruence Q. The dual vector Rt=(R/t)is tangent to this dual curve;

    Rt=Ru1du1dt+Ru2du2dt. (3.1)

    If <Rt,Rt>≠0, then we have a dual unit vector

    R2=RtRt=1P(Ru1du1dt+Ru2du2dt), (3.2)

    where P=p+εp=Ru1du1dt+Ru2du2dt=EG. Hence, the dual arc length of the dual curve R=R(u1(t),u2(t)) is given by

    dS=ds+εds=Pdt. (3.3)

    In order to research the properties of R=R(u1(t),u2(t)), the Blaschke frame relative to R(t) will be defined as the frame of which this line and the central normal R2 to the ruled surface at the central point of R are two edges. The third edge R3 is the central tangent to the ruled surface R(t). Likwise, the frame {R1=R(t), R2=RtRt, R3(t)=R1×R2} is called Blaschke frame. Then we have

    ddS(R1R2R3)=(01010Σ0Σ0)(R1R2R3)=Ω×(R1R2R3), (3.4)

    where

    Ω=ΣR1+R3, (3.5)

    Σ=σ+εσ is the dual geodesic curvature of the dual curve R(t)=R(u1(t),u2(t)). It is easily seen from the last two equations that

    Σ=σ+εσ=det(R,Rt,Rtt)Rt2=det(R,dRdS,d2RdS2). (3.6)

    Referring to the congruence Q, since Rt is tangent of the dual curve R(t), then Eq (3.2) rewritten as

    R2=dS1dSR12+dS2dSR22. (3.7)

    So, we can find a dual angle Θ=ϑ+εϑ such that (Figure 1)

    (R2R3)=(cosΘsinΘsinΘcosΘ)(R12R22), (3.8)
    Figure 1.  R2=cosΘR12+sinΘR22..

    where

    dS2=dS21+dS22,cosΘ=dS1dS=Edu1dS,sinΘ=dS2dS=Gdu2dS.} (3.9)

    By simple verification, it follows that

    dR2dS=R2S1dS1dS+R2S2dS2dS, (3.10)

    we get

    dR2dS=(R12S2+R22S1)sinΘcosΘ+R12S1cos2Θ+R22S2sin2Θ+(R12sinΘ+R22cosΘ)dΘdS. (3.11)

    From Eqs (2.27), (2.29) and (3.8) it follows that

    dR2dS=(Σ1cosΘ+Σ2sinΘ+dΘdS)R3R1. (3.12)

    Thus, from Eqs (3.4) and (3.12), one finds that

    Σ=Σ1cosΘ+Σ2sinΘ+dΘdS. (3.13)

    This formula has the same nature of J. Liouville's formula in surface theory [1,6,9].

    In this subsection we give a geometric characterization of the of J. Liouvile's formula. For this purpose, if R1(t)=R1(t+2π) then R1(t) is called a closed differentiable curve. According to E. Study's map, this corresponds to R1(t) -closed ruled surface belong to the line congruence. Also, let R2 generates a developable ruled surface (torse) along the orthogonal trajectory of the R1(t)-closed ruled surface. From Eq (3.12), this is expressed by

    dR2dS×R1=0Σ1cosΘ+Σ2sinΘ+dΘdS=0. (3.14)

    Then we call the total differential of Θ as the dual angle of pitch of the R1(t)-closed ruled surface. Thus, if it is denoted the dual angle of pitch of the R1(t)-closed ruled surface by the symbol Λ1, then it can be written

    Λ1=λ1εL1:=dΘ, (3.15)

    where λr is the angel of pitch and Lr is the pitch of the R1(t)-closed ruled surface. The pitch and the angle of pitch are well-known real integral invariants of a closed ruled surface [10,14,15,16]. Equations (3.14) and (3.15) shown that

    Λ1=(Σ1cosΘ+Σ2sinΘ)dS. (3.16)

    We found by application of Green's formula, that

    Λ1=(u1(Σ2G)u2(Σ1E))du1du2. (3.17)

    Since the Gaussian curvature of the dual unit sphere can be written by

    1=1EG(u1(Σ2G)u2(Σ1E)). (3.18)

    So that, we obtain as a result the formula

    Λ1=dA1, (3.19)

    where dA1=EGdu1du2 is the dual area on the dual unit sphere enclosed by the closed dual curve R1(t)=R1(t+2π). Hence the following theorem is proved.

    Theorem 1. For a closed ruled surface in the Euclidean 3-space E3. The dual angel of pitch is equal to minus the total dual spherical area of its dual image.

    In fact from Eqs (3.9) and (3.19), we have that

    Λ1:=dS1dS2=(1+ελ1)(1+ελ2)ds1ds2. (3.20)

    If we separate the real and dual parts of Eq (3.20), then we find

    λ1=a1,and   L1=(λ1+λ2)ds1ds2, (3.21)

    where a1 is the element of area on real unit sphere enclosed by the real spherical curve r1(t)=r1(t+2π). More explicitly, we can rewrite Eq (3.16) as

    Λ1=<R12S1,R22>dS1+<R22S1,R12>dS2, (3.22)

    and shown that

    Λ1=Λr1Λr2, (3.23)

    where Λr1, and Λr2 are the dual angel of pitches of the principal ruled surfaces R=R(u1,c2) and R=R(c1,u2), respectively.

    By separating the real and dual parts of Eq (3.9), bearing in mind Eqs (2.25) and (2.26), we get

    λ=λ1cos2ϑ+λ2sin2ϑ,ϑ=(λ2λ12)sin2ϑ.} (3.24)

    These formulas are Hamilton and Mannhiem formulae of surfaces theory in Euclidean 3-space, respectively [1]. The surface described by ϑ in Eq (3.24) is the Plücker conoid. The Plücker conoid is a smooth regular ruled surface sometimes also called the cylindroid [1,6,9].

    The parametric form can also be given in terms of point coordinates. We may choose R1 is coincident with the zaxis of a fixed frame (oxyz), while the position of the dual unit vector R2 is given by angle ϑ and distance ϑ along the positive zaxis. The oriented lines R12 and R22 can be selected in sense of x and yaxes, respectively, as depicted in Figure 2. Clearly, Hamilton's formula shows that the angle ϑ varies from 0 to π2 as λ varies from λ1 to λ2, the principal surfaces being perpendicular to each other. This shows that the oriented lines R12 and R22 are the principal axes and together with the oriented line R1 create the fundamental coordinate system of the Plücker's conoid. Let ζζ denote a point on this surface, it is possible to have the following point coordinates:

    ζζ(ϑ,ϑ,υ)=(0,0,ϑ)+υ(cosϑ,sinϑ,0),  υR. (3.25)
    Figure 2.  Plücker conoid.

    Thus, the zaxis acts as base curve and the circle ϑ(cosϑ,sinϑ,0) as director curve for the parametrization. Using this parametrization, the rulings are clearly visible passing through the zaxis, namely,

    x=υcosϑ,  y=υsinϑ  and   ϑ:=z=(λ2λ1)2sin2ϑ, (3.26)

    which gives us the intersection point of the principal axes R12 and R22 lies at a half of the conoid height ϑ. It can easily be verified by direct computations that

    (x2+y2)z+(λ1λ2)xy=0, (3.27)

    The Eq (3.27) shows that the Plücker's conoid has two integral invariants of the first order, λ1, and λ2. It is assumed by convention that λ1>λ2. The Plücker's conoid depends only on their difference; λ1λ2=1, ϑ[0,2π], υ[1,1] (Figure 2). This surface has two torsal planes π1, π2 and each of which contains one torsal line L as follow: Solving for yx, one obtains a second-order algebraic equation, whose roots are:

    yx=12z[λ2λ1±(λ2λ1)24z2], (3.28)

    The limit points of the Plücker's conoid can be determined by the vanishing of the discriminant of Eq (3.19), which leads to the two extreme positions, that is,

    2z=±(λ2λ1). (3.29)

    According to the value of λ in Eq (3.28), the geometric properties of the Plücker's conoid are discussed as follows:

    A- If the parameter ruled surface R(t)=R(u1(t),u2(t)) is a non-developable surface, i.e. λ0, then there are two real generators passing through the point (0,0,z) only if z<(λ2λ1)/2; for the two limit points z=±(λ2λ1)/2 they coincide with the principal axes R12 and R22.

    B- If the parameter ruled surface R(t)=R(u1(t),u2(t)) is a developable surface, i.e. λ=0, then their two torsal lines L1, L2 are represented by

    L1, L2:yx=tanϑ=±λ1λ2,z=±(λ2λ1)/2, (3.30)

    Thus the two torsal lines L1, and L2 are perpendicular each other, however, only real if λ1λ20. Hence, in this special case, if λ1 and λ2 have the same value, it follows that the Plücker's conoid degenerates into the pencil of lines through "o" in the torsal plane z=0 (and the two isotropic planes through zaxis). Such a line congruence is called an elliptic line congruence. If, however, λ1 and λ2 have opposite signs, the lines L1 and L2 are real and they coincide with the generating lines R12 and R22. Such a line congruence is a hyperbolic line congruence. If either λ1 or λ2 is zero the line congruence is a parabolic line congruence, the lines L1 and L2 both coincide with xaxis; for λ10, and λ2=0 they coincide with yaxis.

    Transition from polar coordinates to Cartesian coordinates could be performed by substituting

    x=cosϑλ,y=sinϑλ, (3.31)

    into Hamilton's formula, one obtain the equation

    D:|λ1|x2+|λ2|y2=1, (3.32)

    of a conic section. As for theory of surfaces, this conic section is the Dupin's indicatrix of the line congruence. We now examine three cases in detail:

    (1) If λ1, and λ2 are both positive, the Dupin's indicatrix is an ellipse has the principal semi-axes are 1λ1, and 1λ2. The lines through the center intersects the ellipse in the points

    x=±cosϑλ,y=±sinϑλ. (3.33)

    The distance intercepted by the ellipse on the line yx=tanϑ is (See Figure 3):

    x2+y2=1λ. (3.34)
    Figure 3.  Dupin's indicatrix with λ1 and λ2 are both positive.

    (2) If λ1, and λ2 have opposite signs the Dupin's indicatrix is set of conjugate hyperbolas

    D:λ1x2λ2y2=±1, (3.35)

    shown in Figure 4. The two asymptotic directions of the hyperbolas represent the torsal lines at which λ=0.

    Figure 4.  Dupin's indicatrix with λ1 and λ2 have opposite signs.

    (3) If either λ1 or λ2 is zero the Dupin's indicatrix is a set of parallels lines corresponding to one of the forms

    y2=|1λ2| with   λ1=0,or   x2=|1λ1| with λ2=0.

    The parametric equation of the dual unit sphere K={RD3R21+R22+R23=1}, may be given by the equations

    R1=cosΘsinΦ,R2=sinΘsinΦ,R3=cosΦ, (3.36)

    where Θ=ϑ+εϑ, and Φ=φ+εφ are dual angles with πϑπ, and π2φπ2. Separating Eq (3.36) into real and dual parts we obtain

    r1=cosϑsinφ, r2=sinϑsinφ,r3=cosφ, (3.37)

    and

    r1=ϑsinϑsinφ+φcosφcosϑ,r2=ϑcosϑsinφ+φcosφsinϑ,r3=φsinφ.} (3.38)

    These coordinates represent the four parameter family of lines in E3. To form line congruence, which describes a two-real parameter motion on the dual unit sphere, we may assume that ϑ=ϑ(ϑ,φ), and φ=φ(ϑ,φ). Thus, the line congruence is given by

    R(ϑ,φ)=(cosϑsinφ,sinϑsinφ,cosφ)+ε[ϑ(sinϑsinφ,cosϑsinφ,0)+φ(cosϑcosφ,sinϑcosφ,sinφ)]. (3.39)

    According to Eqs (2.15) and (2.23), we obtain that

    e=sin2φ,f=0,g=1,e=2(ϑϑsinφ+φcosφ)sinφ,f=(ϑφsin2φ+φϑ),g=2φφ.} (3.40)

    Thus,

    λ=(ϑϑsinφ+φcosφ)sinφdϑ2+(ϑφsin2φ+φϑ)dϑdφ+φφdφ2sin2φdϑ2+dφ2. (3.41)

    In order to identify the principal ruled surfaces of the line congruence, from Eq (3.40), we have

    F=f+εf=0φ=cosϑ, and ϑ=sinϑcotφ, (3.42)

    which yields λ=e=g=f=0. Therefore, the principal ruled surfaces are developable ruled surfaces of the line congruence.

    Now we may calculate the equation of the developable ruled surfaces of the congruence R=R(ϑ,φ) in terms of the Pl ücker coordinates as follows: Since y×r=r we have the system of linear equations in y1, y2 and y3 (yis are the coordinates of y)

    y2cosφy3sinϑsinφ=cosφ,y1cosφ+y3cosϑsinφ=0,y1sinϑsinφy2cosϑsinφ=cosϑsinφ.} (3.43)

    The matrix of coefficients of unknowns y, y2 and y3 is the skew-symmetric matrix

    (    0   cosφsinϑsinφcosφ      0cosϑsinφsinϑsinφcosϑsinφ          0),

    and therefore its rank is 2 with φ0, and ϑ0. Also the rank of the augmented matrix

    (    0    cosφsinϑsinφcosφcosφ      0cosϑsinφ0sinϑsinφcosϑsinφ          0cosϑsinφ),

    is 2. Hence this system has infinite solutions given by

    y1=y3tanφcosϑ,y2=y3tanφsinϑ+1,y3=y3(ϑ,φ). (3.44)

    Since y3(ϑ,φ) can be chosen arbitrarily, then we may take y3ϑ=0. In this case, Eq (3.44) reduces to

    y1(ϑ,φ)=cosϑsinϑ,y2(ϑ,φ)=cos2ϑ,,y3(ϑ,φ)=sinϑcotφ. (3.45)

    According to Eqs (3.37) and (3.45) we obtain

    Y(ϑ,φ,v)=(cosϑsinϑ+vcosϑsinφ,cos2ϑ+vsinϑsinφ,sinϑcotφ+vcosφ).

    It is very clear that if the functions ϑ, and φ are given, then the following developable can be determined. The developable of the congruence is obtained for ϑ(t)=φ(t)=tR, ϑ=const, and φ=const, see Figure 5.

    Figure 5.  Developable surface of the congruence.

    This work mainly deals with the dual representation of line congruence and explains the resemblance between theory of surfaces and theory of line congruences. In terms of this, several some new and well-known formulae of line congruence in the Euclidean 3-space have been introduced. Furthermore, we determine kinematic-geometry of the Plucker conoid and its characterization. In addition, the degenerated cases of the Plücker conoid are discussed according to the Dupin's indicatrix having specific trajectories. We hope that this work will lead to a wider usage in the differential line geometry and rational design of space mechanisms.

    The authors declare that they have no conflict of interest.



    Conflict of interest



    The use of L. enzymogenes B25 as a biocontrol agent is subjected to a patent application. BLG, CF, IH, LPE, MA, ODP, and SMS were employed by Futureco Bioscience S.A.

    Author contributions:



    BLG and SMS conceived the work, contributed to the design and interpretation of the results and did the main writing of the article. CF, IH, LPE, and MA partially wrote specific sections. IH, LPE, MA, and ODP performed the experiments presented in the work and analyzed the data. CF supervised the work and revised the manuscript. All the authors have read the manuscript and agreed to its content.

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