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Review

Phospholipid synthetic and turnover pathways elicited upon exposure to different xenobiotics

  • Received: 25 March 2020 Accepted: 15 June 2020 Published: 18 June 2020
  • Phospholipids, neutral lipids and glycolipids metabolism take the place during cell responses to different stimuli. Phospholipase and acyl glycerol lipase activities have been demonstrated to release second messengers that trigger cascade responses. Studies on lipid droplets structure, formation and interaction with other organelles have recently been described in prokaryotes and eukaryotes. Remodeling of membrane phospholipids has also been reported. NMR studies performed on synthetic membranes allowed to postulate membrane lipids polymorphism and to explain how processes like cell division, endocytosis and exocytosis can take place, due to special arrangements of the membranes. Studies from our research group on the pesticide dieldrin and Cu2+ effects through exposure of amphibian oocytes and embryos to sub-lethal and acclimation concentrations followed by toxic concentration challenges respectively, are discussed. Membrane phospholipids structure alterations allowing stabilization of bilayer arrangements were found for both stressors in cell and tissues. Metallothionein induction response that prevents oxidative stress was also found in acclimated embryo tissues. Probable connections between enzyme activities taking place on lysophospholipid and triacylglycerol substrates present in lipid droplets as well as phospholipid trafficking leading to modifications on membrane phospholipid ratios are discussed. Current new evidences in agreement with our findings allow us to suggest that our previously published results of dieldrin effects on amphibian cells correspond to joint coordinated activities, which probably had involved various metabolic pathways, in line with the acclimation experiment results and discussion. Independently of the cascade responses each toxicant is able to elicit, lipids play an important role in cell responses, both through rapid turnover and final stabilization of membrane bilayers.

    Citation: Teresa M. Fonovich. Phospholipid synthetic and turnover pathways elicited upon exposure to different xenobiotics[J]. AIMS Molecular Science, 2020, 7(3): 211-228. doi: 10.3934/molsci.2020010

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  • Phospholipids, neutral lipids and glycolipids metabolism take the place during cell responses to different stimuli. Phospholipase and acyl glycerol lipase activities have been demonstrated to release second messengers that trigger cascade responses. Studies on lipid droplets structure, formation and interaction with other organelles have recently been described in prokaryotes and eukaryotes. Remodeling of membrane phospholipids has also been reported. NMR studies performed on synthetic membranes allowed to postulate membrane lipids polymorphism and to explain how processes like cell division, endocytosis and exocytosis can take place, due to special arrangements of the membranes. Studies from our research group on the pesticide dieldrin and Cu2+ effects through exposure of amphibian oocytes and embryos to sub-lethal and acclimation concentrations followed by toxic concentration challenges respectively, are discussed. Membrane phospholipids structure alterations allowing stabilization of bilayer arrangements were found for both stressors in cell and tissues. Metallothionein induction response that prevents oxidative stress was also found in acclimated embryo tissues. Probable connections between enzyme activities taking place on lysophospholipid and triacylglycerol substrates present in lipid droplets as well as phospholipid trafficking leading to modifications on membrane phospholipid ratios are discussed. Current new evidences in agreement with our findings allow us to suggest that our previously published results of dieldrin effects on amphibian cells correspond to joint coordinated activities, which probably had involved various metabolic pathways, in line with the acclimation experiment results and discussion. Independently of the cascade responses each toxicant is able to elicit, lipids play an important role in cell responses, both through rapid turnover and final stabilization of membrane bilayers.


    Let p be a prime, f be a polynomial with k variable and Fp=Z/(p) be the finite field, where Z is the integer ring, and let

    N(f;p)=#{(x1,x2,,xk)Fkp|f(x1,x2,,xk)=0}.

    Many scholars studied the exact formula (including upper bound and lower bound) for N(f;p) for many years, it is one of the main topics in the finite field theory, the most elementary upper bounds was given as follows (see [14])

    N(f;p)pk1degf.

    Let ordp denote the p-adic additive valuation normalized such that ordpp=1. The famous Chevalley-Warning theorem shows that ordpN(f;p)>0 if n>degf. Let [x] denote the least integer x and e denote the extension degree of Fq/Fp. Ax (see [2]) showed that

    ordpN(f;q)e[ndegfdegf].

    In 1977, S. Chowla et al. (see [7]) investigated a problem about the number of solutions of a equation in finite field Fp as follow,

    x31+x32++x3k0,

    where p is a prime with p1mod 3 and xiFp, 1ik.

    Let Mk denotes the number of solutions of the above equation. They proved that

    M3=p2+d(p1),M4=p2+6(p2p),s=1Msxs=x1px+x2(p1)(2+dx)13px2pdx3,

    where d is uniquely determined by 4p=d2+27y2 and d1 mod 3.

    Myerson [12] extended the result in [2] to the field Fq and first studied the following equation over Fq,

    x31+x32++x3k0.

    Recently J. Zhao et al. (see [17]) investigated the following equations over field Fp,

       f1=x41+x42+x43,f2=x41+x42+x43+x44.

    And they give exact value of N(f1;p) and N(f2;p). For more general problem about this issue interested reader can see [6,9,10,11].

    In this paper, let A(k,p) denotes the number of solutions of the following equation in Fp,

    x61+x62++x6k0,

    where p is a prime with p1mod 3 and xiFp, 1ik, and for simplicity, in the rest of this paper, we assume there exists an integer z such that z32mod p, we use analytic methods to give a recurrence formula for the number of solutions of the above equation. And our method is based on the properties of Gauss sum. It is worth noting that we used a novel method to simplify the steps and avoid a lot of complicated calculations. We proved the following:

    Theorem 1. For any positive integer k1, we have the recurrence formula

    A(k+6,p)=5pA(k+4,p)+10dpA(k+3,p)+(46p2+5d2p+dp)A(k+2,p)+(2p2+120dp2+3d3p+d2p+dp)A(k+1,p)+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)A(k,p)+pk+5pk+4(10dp+2d2)pk+364pk+5/2(429+121d+5d2)pk+22d2pk+3/2(3d3+130d2+12d+6)pk+1d4pk,

    with the initial condition

    A(1,p)=1,A(2,p)=4(p1)+p,A(3,p)=10d(p1)+p2,A(4,p)=56p(p1)+10d2(p1)+p3,A(5,p)=188dp(p1)+5d3(p1)+16dC(p)(p1)+p4,A(6,p)=p5+1400p2(p1)+(388d2+8d576)p(p1)+d2pd2,

    where d is uniquely determined by 4p=d2+27y2 and d1 mod 3, and C(p)=pa=1ep(a3).

    Remark. Our method is suitable to calculus the number of solutions of the following equation in Fp,

    xt1+xt2++xtk0,

    where p satisfied a certain congruence conditions, and t is any nature number.

    Our Theorem 2 can be deduced from Theorem 1 and the theory of the Difference equations.

    Theorem 2. Let ti (1ik) be the real root of the below equation with multiplicity si (1ik) respectively, and ρje±iwj (1jh) be the complex root of the below equation with multiplicity rj (1jh) respectively,

    x6=5px4+10dpx3+(46p2+5d2p+dp)x2+(2p2+120dp2+3d3p+d2p+dp)x+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p).

    We have

    A(n,p)=ki=1sia=1Ciansiatni+hj=1rjb=1Djbnrjbρnjcosnwj+hj=1rjb=1Ejbnrjbρnjsinnwj,

    where Cia,Djb,Ejb, are determined by

    A(6,p)=ki=1sia=1Cia6siat6i+hj=1rjb=1Djb6rjbρ6jcos6wj+hj=1rjb=1Ejb6rjbρ6jsin6wj,A(5,p)=ki=1sia=1Cia5siat5i+hj=1rjb=1Djb5rjbρ5jcos5wj+hj=1rjb=1Ejb5rjbρ5jsin5wj,A(4,p)=ki=1sia=1Cia4siat4i+hj=1rjb=1Djb4rjbρ4jcos4wj+hj=1rjb=1Ejb4rjbρ4jsin4wj,A(3,p)=ki=1sia=1Cia3siat3i+hj=1rjb=1Djb3rjbρ3jcos3wj+hj=1rjb=1Ejb3rjbρ3jsin3wj,A(2,p)=ki=1sia=1Cia2siat2i+hj=1rjb=1Djb2rjbρ2jcos2wj+hj=1rjb=1Ejb2rjbρ2jsin2wj,A(1,p)=ki=1sia=1Ciati+hj=1rjb=1Djbρjcoswj+hj=1rjb=1Ejbρjsinwj. (1.1)

    Before we prove these lemmas, we give some notations, χ2 denotes the second-order character of Fp, χ denotes the third-order character of Fp, ψ denotes the sixth order character of Fp.

    ep(x)=e2πixp,τ(χ)=pm=1χ(m)ep(m),G(χ,m)=pa=1χ(a)ep(am).

    We call G(χ,m) the Gauss sum, and we have the following:

    G(χ,m)=τ(χ)¯χ(m),(m,p)=1. (2.1)

    And also we have

    |τ(χ)|=p, (2.2)

    where χ is a primitive character of Fp. And let G(m,6;p)=p1a=0ep(ma6). For the property of the exponential sum and the general Gauss sum, interested readers can see [1,4,5,8,13,15].

    Lemma 1. Let p be a prime with p1mod 3. Then for any third-order character χ of Fp, we have the identity

    τ3(χ)+τ3(¯χ)=dp,

    where d is uniquely determined by 4p=d2+27y2 and d1mod 3.

    Proof. For the proof of this lemma see [3].

    Lemma 2. Let χ be a third-order character of Fp with p1mod 3, and C(p)=τ(χ)+τ(¯χ), then C(p)=pa=1ep(a3).

    Proof.

    A=τ(χ)+τ(¯χ)=pa=1(1+χ(a)+¯χ(a))e(ap)=pa=1e(a3p).

    Lemma 3. Let p1mod 6, 2z3mod p for some z, and let χ be a third-order character of Fp, ψ be a sixth-order character of Fp, then we have the identity

    τ(ψ)=τ2(χ)p.

    Proof. This is Lemma 3 in [16].

    Lemma 4. As the definition above, we have the identity

    G(m,6;p)=pχ2(m)+¯x2pψ(m)+x2p¯ψ(m)+¯xχ(m)+x¯χ(m),

    where (m,p)=1 and x=τ(χ).

    Proof. Firstly we have the identity

    1+χ2(m)+χ(m)+¯χ(m)+ψ(m)+¯ψ(m)={6, if  ma6 mod p;0,otherwise.

    So we have

    G(m,6;p)=p1a=0(1+χ2(a)+χ(a)+¯χ(a)+ψ(a)+¯ψ(a))ep(ma)=G(χ2,m)+G(ψ,m)+G(¯ψ,m)+G(χ,m)+G(¯χ,m)

    By (2.1) and Lemma 3, we have

    G(m,6;p)=τ(χ2)χ2(m)+τ(¯ψ)ψ(m)+τ(ψ)¯ψ(m)+τ(¯χ)χ(m)+τ(χ)¯χ(m)=pχ2(m)+¯x2pψ(m)+x2p¯ψ(m)+¯xχ(m)+x¯χ(m). (2.3)

    By (2.3), we complete the proof of our lemma.

    Next we let,

    Gn(m,6;p)=an+bnχ2(m)+cnψ(m)+dn¯ψ(m)+enχ(m)+fn¯χ(m). (2.4)

    Then we have following Lemma 5.

    Lemma 5. Let an,bn,cn,dn,en,fn are defined as above, then we have that an,bn,cn,dn,en,fn are uniquely determined by n, where n1.

    Proof. By the orthogonality of characters of Fp, we have

    p1a=1χ(a)={p1, if χ=χ0;0,otherwise. (2.5)

    By (2.4) and (2.5) we have

    p1m=1Gn(m,6;p)=(p1)an+bnp1m=1χ2(m)+cnp1m=1ψ(m)+dnp1m=1¯ψ(m)+enp1m=1χ(m)+fnp1m=1¯χ(m)=(p1)an.

    So we have

    an=1p1p1m=1Gn(m,6;p). (2.6)

    By the same method, we have

    bn=1p1p1m=1χ2(m)Gn(m,6;p),cn=1p1p1m=1¯ψ(m)Gn(m,6;p),dn=1p1p1m=1ψ(m)Gn(m,6;p),en=1p1p1m=1¯χ(m)Gn(m,6;p),fn=1p1p1m=1χ(m)Gn(m,6;p).

    So now it is easy to see the conclusion of the lemma.

    Lemma 6. The sequences {an}, {bn}, {cn}, {dn}, {en}, {fn} are defined above, then they satisfied the following recurrence formulae (n0):

    an+1=pbn+¯x2pdn+x2pcn+xen+¯xfn, (2.7)
    bn+1=pan+¯x2pen+x2pfn+xdn+¯xcn, (2.8)
    cn+1=pfn+¯x2pan+x2pen+xbn+¯xdn, (2.9)
    dn+1=pen+¯x2pfn+x2pan+xcn+¯xbn, (2.10)
    en+1=pdn+¯x2pcn+x2pbn+xfn+¯xan, (2.11)
    fn+1=pcn+¯x2pbn+x2pdn+xan+¯xen, (2.12)

    with the initial condition

    a0=1,b0=c0=d0=e0=f0=0.

    Proof. We only prove (2.7), the rest can be proved in the same way. By Lemma 5, we know an is unique determined by n. We can compare the coefficient of the equation

    Gn+1(m,6;p)=Gn(m,6;p)G(m,6;p).

    We have

    an+1=pbn+¯x2pdn+x2pcn+xen+¯xfn.

    So we complete the proof of the lemma.

    Lemma 7. Let an is defined as above, then we have

    a0=1,a1=0,a2=5p,a3=10dp,a4=56p2+10d2p,a5=188dp2+5d3p+16dpC(p).

    Proof. By Lemma 4 and after some elementary calculations we have

    G2(m,6;p)=5p+2dp1/2χ2(m)+4p1/2xψ(m)+4p1/2¯x¯ψ(m)+(p1¯x4+3x2)χ(m)+(p1x4+3¯x2)¯χ(m),G3(m,6;p)=10dp+(16p3/2+dp1/2)χ2(m)+(15p¯x+2dx2+p1x5)χ(m)+(15px+2d¯x2+p1¯x5)¯χ(m)+(4p1/2x4+12p1/2¯x2+2dp1/2x)ψ(m)+(4p1/2¯x4+12p1/2x2+2dp1/2¯x)¯ψ(m),G4(m,6;p)=60p2+9d2p+dp+48dp3/2χ2(m)+(p2x8+17¯x4+46px2+16dp)χ(m)+(p2¯x8+17x4+46p¯x2+16dp)¯χ(m)+(56p3/2x+4dp1/2x4+12dp1/2¯x2+8p1/2¯x5)ψ(m)+(56p3/2¯x+4dp1/2¯x4+12dp1/2x2+8p1/2x5)¯ψ(m),G5(m,6;p)=188dp2+5d3p+16dpC(p)+(52d2p3/2+208p5/2+16dp1/2(x2+¯x2))χ2(m)+(p2/5x10+p3/2¯x8+4dp1/2¯x5+71p1/2x4+(46p3/2+16p1/2)x2+(129p3/2+10d2p1/2)¯x2+60dp3/2x+16dp3/2)ψ(m)+(p2/5¯x10+p3/2x8+4dp1/2x5+71p1/2¯x4+(46p3/2+16p1/2)¯x2+(129p3/2+10d2p1/2)x2+60dp3/2¯x+16dp3/2)¯ψ(m)+(8p1¯x7+p1x7+25x5+52dpx2+(28dp+46p2)x+16d¯x4+112p2¯x)χ(m)+(8p1x7+p1¯x7+25¯x5+52dp¯x2+(28dp+46p2)¯x+16dx4+112p2x)¯χ(m),

    and comparing the above formulae with (2.6), we have

    a0=1,a1=0,a2=5p,a3=10dp,a4=60p2+9d2p+dp,a5=188dp2+5d3p+16dpC(p).

    Lemma 8. Let an, bn, cn, dn, en, fn are defined as above, then we have

    a6=5pa4+10dpa3+(46p2+5d2p+dp)a2+(2p2+120dp2+3d3p+d2p+dp)a1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)a0b6=5pb4+10dpb3+(46p2+5d2p+dp)b2+(2p2+120dp2+3d3p+d2p+dp)b1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)b0c6=5pc4+10dpc3+(46p2+5d2p+dp)c2+(2p2+120dp2+3d3p+d2p+dp)c1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)c0d6=5pd4+10dpd3+(46p2+5d2p+dp)d2+(2p2+120dp2+3d3p+d2p+dp)d1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)d0e6=5pe4+10dpe3+(46p2+5d2p+dp)e2+(2p2+120dp2+3d3p+d2p+dp)e1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)e0f6=5pf4+10dpf3+(46p2+5d2p+dp)f2+(2p2+120dp2+3d3p+d2p+dp)f1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)f0

    Proof. We only proof the first formula, the rest can be proof in the same way. By Lemma 6, we have

    a6=pb5+¯x2pd5+x2pc5+xe5+¯xf5=5pa4+2dp1/2b4+4p1/2¯xc4+4p1/2xd4+(3¯x2+p1x4)e4+(3x2+p1¯x4)f4=5pa4+10dpa3+(d2p1/2+12p3/2)b3+(2dp1/2¯x+8p1/2x2+p1/2¯x4)c3+(2dp1/2x+8p1/2¯x2+p1/2x4)d3+(11px+¯x2+p1¯x5)e3+(11p¯x+x2+p1x5)f3=5pa4+10dpa3+(46p2+5d2p+dp)a2+(25dp3/2+2p3/2)b2+(p3/2¯x7+2p1/2x5+p1/2¯x4+42p3/2¯x+2dp1/2x2+(d2+1)p1/2¯x)c2+(p3/2x7+2p1/2¯x5+p1/2x4+42p3/2x+2dp1/2¯x2+(d2+1)p1/2x)d2+(10x4+(32p+d2)¯x2+(4dp+p)x)e2+(10¯x4+(32p+d2)x2+(4dp+p)¯x)f2=5pa4+10dpa3+(46p2+5d2p+dp)a2+(2p2+120dp2+3d3p+d2p+dp)a1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)a0.

    So we complete the proof of this lemma.

    Lemma 9. Let an is defined as above, then for any integer n0, we have

    an+6=5pan+4+10dpan+3+(46p2+5d2p+dp)an+2+(2p2+120dp2+3d3p+d2p+dp)an+1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)an.

    Proof. By (2.4) and Lemma 8, we have

    G6(m,6;p)=5pG4(m,6;p)+10dpG3(m,6;p)+(46p2+5d2p+dp)G2(m,6;p)+(2p2+120dp2+3d3p+d2p+dp)G(m,6;p)+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p).

    We multiple Gn(m,6;p) to the both side of the above formula, we have

    Gn+6(m,6;p)=5pGn+4(m,6;p)+10dpGn+3(m,6;p)+(46p2+5d2p+dp)Gn+2(m,6;p)+(2p2+120dp2+3d3p+d2p+dp)Gn+1(m,6;p)+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)Gn(m,6;p).

    By Lemma 5, we can compare the coefficient of the above equation, we have

    an+6=5pan+4+10dpan+3+(46p2+5d2p+dp)an+2+(2p2+120dp2+3d3p+d2p+dp)an+1+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)an.

    In the formula below, we always let k1. By the following formula,

    p1a=0ep(ma)={p, if  pm;0, otherwise, 

    we have

    A(k,p)=1pp1m=0p1x1=0,x2=0,,xk=0ep(m(x61+x62++x6k))=1pp1m=0Gk(m,6;p). (3.1)

    By (8), we have

    A(k,p)=1pp1m=0Gk(m,6;p)=1p(p1m=1Gk(m,6;p)+pk)=1p((p1)ak+pk)=p1pak+pk1. (3.2)

    So by Lemma 9, we have

    A(k+6,p)pk+5=5p(A(k+4,p)pk+3)+10dp(A(k+3,p)pk+2)+(46p2+5d2p+dp)(A(k+2,p)pk+1)+(2p2+120dp2+3d3p+d2p+dp)(A(k+1,p)pk)+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)(A(k,p)pk1).

    So we have

    A(k+6,p)=5pA(k+4,p)+10dpA(k+3,p)+(46p2+5d2p+dp)A(k+2,p)+(2p2+120dp2+3d3p+d2p+dp)A(k+1,p)+(4p5+2d2p4+64p7/2+381p3+2d2p5/2+(129d2+11d+6)p2+d4p)A(k,p)+pk+5pk+4(10dp+2d2)pk+364pk+5/2(429+121d+5d2)pk+22d2pk+3/2(3d3+130d2+12d+6)pk+1d4pk.

    And by Lemma 7 and (3.2), we have the initial conditions

    A(1,p)=1,A(2,p)=4(p1)+p,A(3,p)=10d(p1)+p2,A(4,p)=56p(p1)+10d2(p1)+p3,A(5,p)=188dp(p1)+5d3(p1)+16dC(p)(p1)+p4.A(6,p)=p5+1400p2(p1)+(388d2+8d576)p(p1)+d2pd2.

    So we complete the proof of the theorem.

    The main purpose of this paper is using analytic methods to give a recurrence formula of the number of solutions of an equation over finite field. And we give an expression of the number of solutions of the above equation by the root of sixth degree polynomial. We use analytic methods to give a recurrence formula for the number of solutions of the above equation. And our method is based on the properties of the Gauss sum. It is worth noting that we used a novel method to simplify the steps and avoid complicated calculations.

    The author thanks to referees for very important recommendations and warnings which improved the paper.

    The author declares that there is no competing interest.


    Acknowledgments



    This work was supported in part by the Science, Technology and Innovation Ministry (under grant FONCyT PICT 2017-0706) and by the School of Science and Technology from National University of San Martín. Figures 1 and 2 were illustrated by Silvina Mariel Schroeder. The author appreciates Cristina Pérez Coll kindly provision of embryo photograph shown in Figure 2. Photograph shown in Figure 1 belongs to the author.

    Conflict of interest



    The author declares no conflict of interest for the contributions in this manuscript.

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