Somitogenesis is the process by means of which a tissue known as presomitic mesoderm (PSM) is segmented in blocks of cells, called somites, along the anterior-posterior axis of the developing embryo in segmented animals. In vertebrates, somites give rise to axial skeleton, cartilage, tendons, skeletal muscle, and dermis. Somite formation occurs periodically, and this periodicity is driven by a genetic oscillator that operates within PSM cells and is known as the segmentation clock. The correct synchronization of the segmentation clock among PSM cells is essential for somitogenesis to develop normally. When synchronization is disrupted, somites form irregularly and, in consequence, the tissues that originate from them show clear malformations. In this work, based in a model for zebrafish segmentation clock, we investigate by means of a mathematical modeling approach, how PSM-cell synchronization is affected by factors like: the size of PSM-cell networks, the amount of cell-to-cell interactions per PSM cell, the strength of these interactions, and the inherent variability among PSM cells. Interestingly we found that very small PSM-cell networks are unable to synchronize. Moreover, the effect of decreasing the strength of interactions among PSM cells is corrected by increasing the network connectivity-level, and a moderated level of variability among cells can have a positive effect on synchronization, specially in large networks.
Citation: Jesús Pantoja-Hernández, Moisés Santillán. Segmentation-clock synchronization in circular-lattice networks of embryonic presomitic-mesoderm cells[J]. AIMS Mathematics, 2021, 6(6): 5817-5836. doi: 10.3934/math.2021344
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Somitogenesis is the process by means of which a tissue known as presomitic mesoderm (PSM) is segmented in blocks of cells, called somites, along the anterior-posterior axis of the developing embryo in segmented animals. In vertebrates, somites give rise to axial skeleton, cartilage, tendons, skeletal muscle, and dermis. Somite formation occurs periodically, and this periodicity is driven by a genetic oscillator that operates within PSM cells and is known as the segmentation clock. The correct synchronization of the segmentation clock among PSM cells is essential for somitogenesis to develop normally. When synchronization is disrupted, somites form irregularly and, in consequence, the tissues that originate from them show clear malformations. In this work, based in a model for zebrafish segmentation clock, we investigate by means of a mathematical modeling approach, how PSM-cell synchronization is affected by factors like: the size of PSM-cell networks, the amount of cell-to-cell interactions per PSM cell, the strength of these interactions, and the inherent variability among PSM cells. Interestingly we found that very small PSM-cell networks are unable to synchronize. Moreover, the effect of decreasing the strength of interactions among PSM cells is corrected by increasing the network connectivity-level, and a moderated level of variability among cells can have a positive effect on synchronization, specially in large networks.
Let q be a power of odd prime. Several researchers have looked into a variety of properties about the primitive roots modulo q. Let g1,g2 represent two primitive roots modulo q, a, b and c represent arbitrary non-zero elements in Fq. Is there some q0 such that for all q>q0, there is always one representation
a=bg1+cg2 ? | (1.1) |
For b=1 and c=−1, Vegh [1] considered a specific form of Eq (1.1), which is known as Vegh's Conjecture, (see [2,§ F9] for further details). Cohen [3] demonstrated Vegh's Conjecture for all q>7.
For b=1 and c=1, Golomb [4] proposed another specific form of Eq (1.1). This was proved by Sun [5] for q>260≈1.15×1018.
Moreover, Cohen et al. [6] studied linear sums of primitive roots and their inverses in finite fields Fq and showed that if q>13, then for arbitrary non-zero a,b∈Fq, there is a pair of primitive elements (g1, g2) of Fq such that both ag1+bg2 and ag−11+bg−12 are primitive.
Let p be an odd prime. Carlitz [7] relied on some results of Davenport and obtained for any k−1 fixed integers c1,c2,…,ck−1 with ci≥1(i=1,2,…,k−1). Let g,g1,…,gk−1 be primitive roots modulo p and Nk denote the number of gmodp such that g1−g=c1,…,gk−1−g=ck−1. Then
Nk∼ϕk(p−1)pk−1 (p→∞). |
More results of the primitive roots distribution can be found in [8,9,10,11].
Lehmer [2,§ F12] proposed the definition of Lehmer number, according to which a is a Lehmer number if and only if a and ˉa have opposite parity, i.e., (2,a+ˉa)=1, where ˉa is the multiplicative inverse of a modulo p. It is simple to demonstrate that there are no Lehmer numbers modulo p when p=3 or 7. Zhang [12] established that if Mp denotes the number of Lehmer numbers modulo p, then
Mp=p−12+O(p12ln2p). |
A Lehmer number that is also a primitive root modulo p will be called a Lehmer primitive root or an LPR. The inverse of an LPR is also an LPR. We assume that p>3 because there is no Lehmer number modulo 3. Wang and Wang [13] investigated the distribution of LPRs involving Golomb's conjecture. Let Gp denote the number of Golomb pairs (a,b) (i.e., a+b≡1(modp)) are LPRs. They showed
Gp=14ϕ2(p−1)p−1+O(ϕ2(p−1)p54⋅4ω(p−1)⋅ln2p). |
Let Np denote the number of LPRs modulo p. For odd integers m≥3, define the positive number Tm by
Tm=2mlnm(m−1)/2∑j=1tan(πjm). |
Cohen and Trudgian [14] improved the result of Wang and Wang [13] and showed
|Np−ϕ(p−1)2|<T2pϕ(p−1)p−12ω(p−1)p12ln2p |
and
|Gp−ϕ2(p−1)4(p−1)2(p−2)|<ϕ2(p−1)4(p−1)2T2p[22ω(p−1)(9ln2p+1)−1]p12, |
where 2π(1+0.548lnp)<Tp<2π(1+1.549lnp).
Specifically, they obtained that for an odd prime p(≠3,7), there exists an LPR modulo p.
Inspired by the results of Cohen and Trudgian [14] and Wang and Wang [13], we mainly studied the distribution of LPRs modulo p related to the Golomb's conjecture in two aspects. On the one hand, we extend Eq (1.1) to the case involving k>1 variables. Let R be set of LPRs modulo p that is a subset of Fp. a1,a2,…,ak,c are non-zero elements in Fp and N(R,p) denotes the number of solutions of the equation
a1g1+a2g2+⋯+akgk=c, g1,g2,…,gk∈R. |
We consider the distribution properties of N(R,p), and obtain the following:
Theorem 1. Let p>3 be an odd prime. Then we have
N(R,p)=ϕk(p−1)2kp+O(ϕk(p−1)p322kω(p−1)ln2kp), |
where the symbol O is dependent on k.
When k=2, we can obtain the number of the Golomb pairs that are LPRs.
On the other hand, we consider the distribution of k consecutive LPRs and generalize it to a more general form.
Let f(x)∈Fp[x]. Define
M(f(x),R,p)=#{x:1≤x≤p−1,f(x+c1),f(x+c2),⋯,f(x+ck)∈R}. |
Then we have:
Theorem 2. Let f(x)∈Fp[x] with degree l≥1. c1,c2,…,ck are distinct elements in Fp. Suppose that one of the following conditions holds:
(i) f(x) is irreducible,
(ii) f(x) has no multiple zero in ˉFp and k=2,
(iii) f(x) has no multiple zero in ˉFp and (4k)l<p.
Then we have
M(f(x),R,p)=12kϕk(p−1)(p−1)k−1+O(ϕk(p−1)pk−122kω(p−1)ln2kp), |
where the symbol O is dependent on k and l.
Take f(x)=x, ck=0 in Theorem 2. Then we can get the number of k consecutive primitive roots x,x+c1,…,x+ck−1 are Lehmer numbers, which is:
Corollary 1. Let p be an odd prime. Then for any 1≤ x≤(p−1) that is an LPR modulo p, we have
M(x,R,p)=12kϕk(p−1)(p−1)k−1+O(ϕk(p−1)pk−122kω(p−1)ln2kp), |
where the symbol O is dependent on k.
When k=1,2, we can easily deduce the Theorem 1 and Theorem 6 in Cohen and Trudgian [14], respectively.
Notation: Throughout this paper, Fq denotes a finite field of characteristic p, ˉFq denotes the algebraic closure of Fq, ϕ(n) is reserved for the Euler function, μ(n) is the M¨obius function. We use ω(n) to denote the number of all distinct prime divisors of n. Write ∑χd to denote a sum over all ϕ(d) multiplicative characters χd of order d over Fp, and denote by ∑pn=1′ the summation of 1≤n≤p with (n,p)=1. τ(χ) is the classical Gauss sums associated with character χ mudulo p. f≪g means |f|≤cg with some positive constant c, f=O(g) means f≪g.
To complete the proof of the theorems, we need following several lemmas. The proofs of these lemmas require some basic knowledge of analytic number theory, which can be found in [15].
Lemma 1. Let p be an odd prime. Then for any integer a coprime to p (i.e., (a,p)=1), we have the identity
ϕ(p−1)p−1∑d∣p−1μ(d)ϕ(d)∑χdχd(a)={1, if a is a primitive root mod p;0, if a is not a primitive root mod p. |
Proof. See Proposition 2.2 of Narkiewicz [16].
Lemma 2. Let p be an odd prime, χ be a nonprincipal multiplicative character modulo p of order d. Suppose g(x)∈Fp[x] has precisely m distinct ones among its zeros, and suppose that g(x) is not the constant multiple of a d-th power over Fq. Then
|∑x∈Fpχ(g(x))|≤(m−1)⋅p12. |
Proof. See Theorem 2C in Chapter 2 of Schmidt [17].
Lemma 3. Let Fq be a finite field of characteristic p, ψ be a nontrivial additive character and χ be a nonprincipal multiplicative character on Fq of order d. For two rational functions f(x),g(x)∈Fq[x], define K(ψ,f;χ,g)=∑x∈Fq∖Sχ(g(x))ψ(f(x)), where S denotes the set of poles of f(x) and g(x). Suppose the following conditions hold:
(i) g(x) is not the constant multiple of a d-th power over Fq.
(ii) f(x) is not of the form (h(x))p−h(x) with a rational function h(x) over Fq.
Then we have
|K(ψ,f;χ,g)|≤(deg(f)+m−1)√q, |
where m is the number of distinct roots and (noninfinite) poles of g(x) in Fq.
Proof. See Theorem 2G in Chapter 2 of Schmidt [17].
Lemma 4. Let p be an odd prime. Let c1,⋯,ck be distinct elements in Fp. Assume that f(x)∈Fp[x] with deg(f)=l. Define the polynomial
h(x)=f(x+c1)⋯f(x+ck). |
Suppose one of the following conditions holds:
(i) f(x) is irreducible,
(ii) f(x) has no multiple zero in ˉFp and k=2,
(iii) f(x) has no multiple zero in ˉFp and (4k)l<p.
Then h(x) has at least one simple root in ˉFp.
Proof. Suppose that f(x) is irreducible. Then f(x+c1),⋯,f(x+ck) are distinct irreducible polynomials, and h(x) has at least k simple roots in ˉFp. The cases of (ii) and (iii) can be proved by Theorem 2 and Lemma 2 of [18], for k=2 or (4k)l<p, (l,k,p) is "admissible triple, " then f(x+c1)⋯f(x+ck) has at least one simple root.
Lemma 5. Let p be an odd prime, m1,…,mk,n1,…,nk be integers with (m1⋯mkn1⋯nk,p)=1, and polynomials g(x),f1(x),…,fk(x)∈Fp[x]. Let χ be a Dirichlet character modulo p of order d. Define
K(χ,g,f1,⋯,fk;p)=p∑x=1(f1(x)⋯fk(x),p)=1χ(g(x))e(m1f1(x)+⋯+mkfk(x)+n1¯f1(x)+⋯+nk¯fk(x)p). |
Suppose the following conditions hold:
(i) g(x) can not be the constant multiple of a d-th power over Fp.
(ii) F(x)=f1(x)⋯fk(x) has at least one simple root in ˉFp.
Then we have
|K(χ,g,f1,⋯,fk;p)|≤(max(deg(f1),⋯,deg(fk))+l)√p, |
where e(x)=e2πix and l is the number of distinct roots of g(x) in ˉFp.
Proof. It is clear that
m1f1(x)+⋯+mkfk(x)+n1¯f1(x)+⋯+nk¯fk(x)=F(x)(m1f1(x)+⋯+mkfk(x))+n1F(x)f1(x)+⋯+nkF(x)fk(x)F(x):=G(x)F(x). |
Condition (i) is the same as Lemma 3. So our goal is to prove the rational function G(x)/F(x) satisfies condition (ii) in Lemma 3 if F(x) has a simple root in ˉFp. Assume that there are polynomials K(x),L(x)∈Fp[x] with (K(x),L(x))=1 such that
G(x)F(x)=(K(x)L(x))p−(K(x)L(x)). |
Then we have
G(x)L(x)p=(K(x)p−K(x)L(x)p−1)F(x). | (2.1) |
Since F(x)=f1(x)⋯fk(x) has at least one simple root in ˉFp, then there exists an irreducible polynomial w(x)∈Fp[x] such that w(x)∣F(x) and w(x)2∤F(x). Assume that w(x)∣f1(x), then we have
w(x)∤F(x)f1(x), w(x)∣F(x)fi(x)(i=2,⋯,k). |
Hence, from Eq (2.1)
w(x)∤G(x)⟹w(x)∣L(x)p⟹w(x)∣L(x) |
w(x)2∣L(x)p−1⟹w(x)2∣K(x)pF(x)⟹w(x)∣K(x), |
which contradicts to (K(x),L(x))=1. Therefore, from Lemma 3 we get
|K(χ,g,f1,⋯,fk;p)|≤(max(deg(f1),⋯,deg(fk))+l)√p, |
where l is the number of distinct roots of g(x) in ˉFp.
Lemma 6. Let χ be a primitive character modulo p, χdi be character modulo p of order di. There exist some 1≤si≤di with (si,di)=1, i=1,2,…,k. Then we have
∑χd1⋯∑χdkχd1(f(x+c1))⋯χdk(f(x+ck))=d1∑s1=1 ′⋯dk∑sk=1 ′χ((f(x+c1))s1(p−1)d1⋯(f(x+ck))sk(p−1)dk). |
Proof. From the definition of the Dirichlet character modulo p, we can get
∑χd1⋯∑χdkχd1(f(x+c1))⋯χdk(f(x+ck))=d1∑s1=1 ′⋯dk∑sk=1 ′e(s1⋅ind(f(x+c1))d1)⋯e(sk⋅ind(f(x+ck))dk)=d1∑s1=1 ′⋯dk∑sk=1 ′e(s1(p−1)d1⋅ind(f(x+c1))+⋯+sk(p−1)dk⋅ind(f(x+ck))p−1)=d1∑s1=1 ′⋯dk∑sk=1 ′e(ind(f(x+c1))s1(p−1)d1+⋯+ind(f(x+ck))sk(p−1)dkp−1)=d1∑s1=1 ′⋯dk∑sk=1 ′e(ind((f(x+c1))s1(p−1)d1⋯(f(x+ck))sk(p−1)dk)p−1)=d1∑s1=1 ′⋯dk∑sk=1 ′χ(f(x+c1))s1(p−1)d1⋯(f(x+ck))sk(p−1)dk), |
where ind(a) denotes an index of a with base g of modulo p, and g is a positive primitive root of modulo p.
Firstly, we prove the Theorem 1. Let p be an odd prime, k be any fixed positive integer. Then for any k different integers a1, a2,…,ak∈Fp, from Lemma 1 and the definition of Lehmer number we have
N(R,p)=1pp−1∑b=0p−1∑g1=1p−1∑g2=1⋯p−1∑gk=1g1,g2,…,gk∈Re(b(a1g1+⋯+akgk−c)p)=1pϕk(p−1)2k(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip−1∑gi=1χdi(gi)(1−(−1)gi+¯gi))⋅p−1∑b=0e(b(a1g1+⋯+akgk−c)p)=1pϕk(p−1)2k(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip−1∑gi=1χdi(gi))p−1∑b=0e(b(a1g1+⋯+akgk−c)p)+1pϕk(p−1)2k(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip−1∑gi=1χdi(gi))k∑t=1(−1)tk∑i1=1k∑i2=1⋯k∑it=1i1<i2<⋯<itli1li2⋯lit⋅p−1∑b=0e(b(a1g1+⋯+akgk−c)p)=A1+A2, | (3.1) |
where li=(−1)gi+¯gi,i=1,2,⋯,k.
A1=1pϕk(p−1)2k(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip−1∑gi=1χdi(gi))p−1∑b=0e(b(a1g1+⋯+akgk−c)p)=1pϕk(p−1)2k(p−1)k[p−1∑g1=1⋯p−1∑gk=1p−1∑b=0e(b(a1g1+⋯+akgk−c)p)+∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk(gk)⋅p−1∑b=0e(b(a1g1+⋯+akgk−c)p)]=1pϕk(p−1)2k(p−1)k[(p−1)k+(−1)k+1+∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)⋅∑χd1⋯∑χdkp−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk(gk)p−1∑b=0e(b(a1g1+⋯+akgk−c)p)]. | (3.2) |
From Eq (3.2), let
A11=∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk(gk)⋅p−1∑b=0e(b(a1g1+a2g2+⋯+akgk−c)p)=∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk(gk)+∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑b=1p−1∑g1=1χd1(g1)e(ba1g1p)⋯p−1∑gk=1χdk(gk)e(bakgkp)e(−bcp)=∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑b=1p−1∑g1=1χd1(g1)e(ba1g1p)⋯p−1∑gk=1χdk(gk)e(bakgkp)e(−bcp). |
Using the properties of Gauss sums we can get
|A11|=|∑d1∣p−1⋯∑dk∣p−1d1⋯dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑b=1p−1∑g1=1χd1(g1)e(ba1g1p)⋯p−1∑gk=1χdk(gk)e(bakgkp)e(−bcp)|=|∑d1∣p−1d1>1⋯∑dk∣p−1dk>1μ(d1)ϕ(d1)⋯μ(dk)ϕ(dk)∑χd1⋯∑χdkp−1∑b=1p−1∑g1=1χd1(g1)e(ba1g1p)⋯p−1∑gk=1χdk(gk)e(bakgkp)e(−bcp)+∑d1∣p−1d1>1⋯∑dk−1∣p−1dk−1>1μ(d1)ϕ(d1)⋯μ(dk−1)ϕ(dk−1)∑χd1⋯∑χdk−1p−1∑b=1p−1∑g1=1χd1(g1)e(ba1g1p)⋯p−1∑gk−1=1χdk−1(gk−1)e(bak−1gk−1p)p−1∑gk=1e(bakgkp)e(−bcp)+⋯+∑d1∣p−1d1>1μ(d1)ϕ(d1)∑χd1p−1∑b=1p−1∑g1=1χd1(g1)e(ba1g1p)p−1∑g2=1e(ba2g2p)⋯p−1∑gk=1e(bakgkp)e(−bcp)|≪2kω(p−1)pk+12, |
where we have used the fact that ∑d|n|μ(d)|=2ω(n).
Hence, Eq (3.2) and the above formulae yield that
A1=ϕk(p−1)2kp+O(ϕk(p−1)pk+122kω(p−1)). | (3.3) |
Then we compute A2 in Eq (3.1). For simplicity, let
Um(u)=p−1∑u=1(−1)ue(−mup), |
noting that
p−1∑u=1(−1)ue(−mup)=1−e(mp)1+e(mp)=isin(πm/p)cos(πm/p), |
p−1∑m=1|sin(πm/p)cos(πm/p)|=Tpplnp. |
Hence,
|p−1∑m=1Um(u)|≤p−1∑m=1|p−1∑u=1(−1)ue(−mup)|=Tpplnp. | (3.4) |
Noting that, if m=0, then ∑p−1u=1(−1)ue(−mup)=∑p−1u=1(−1)u=0, since p is odd. Hence,
li=(−1)gi+¯gi=1pp−1∑mi=0p−1∑ui=1(−1)uie(mi(gi−ui)p)⋅1pp−1∑ni=0p−1∑vi=1(−1)vie(ni(¯gi−vi)p)=1p2p−1∑mi,ni=0e(migi+ni¯gip)p−1∑ui=1(−1)uie(−miuip)p−1∑vi=1(−1)vie(−nivip)=1p2p−1∑mi,ni=1e(migi+ni¯gip)Umi(ui)Uni(vi). | (3.5) |
From the above discussion and Eq (3.1), we can obtain
|A2|=|1pϕk(p−1)2k(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip−1∑gi=1χdi(gi))k∑t=1(−1)tk∑i1=1⋯k∑it=1i1<⋯<itli1⋯lit⋅p−1∑b=0e(b(a1g1+a2g2+⋯+akgk−c)p)|≤1pϕk(p−1)2k(p−1)kk∑t=1(kt)T2tpln2tp∑d1∣p−1⋯∑dk∣p−1|μ(d1)|ϕ(d1)⋯|μ(dk)|ϕ(dk)∑χd1⋯∑χdk|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk(gk)⋅e(m1g1+n1¯g1+⋯+mtgt+nt¯gtp)e(b(a1g1+⋯+akgk−c)p)|=1pϕk(p−1)2k(p−1)kk∑t=1(kt)T2tpln2tp[∑d1∣p−1d1>1⋯∑dk∣p−1dk>1|μ(d1)|ϕ(d1)⋯|μ(dk)|ϕ(dk)∑χd1⋯∑χdk|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk(gk)⋅e(m1g1+n1¯g1+⋯+mtgt+nt¯gtp)e(b(a1g1+⋯+akgk−c)p)|+∑d1∣p−1d1>1⋯∑dk−1∣p−1dk−1>1|μ(d1)|ϕ(d1)⋯|μ(dk−1)|ϕ(dk−1)∑χd1⋯∑χdk−1|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1χd1(g1)⋯χdk−1(gk−1)⋅e(m1g1+n1¯g1+⋯+mtgt+nt¯gtp)e(b(a1g1+⋯+akgk−c)p)|+⋯+∑d1∣p−1d1>1|μ(d1)|ϕ(d1)∑χd1|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1χd1(g1)⋅e(m1g1+n1¯g1+⋯+mtgt+nt¯gtp)e(b(a1g1+⋯+akgk−c)p)|+|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1e(m1g1+n1¯g1+⋯+mtgt+nt¯gtp)⋅e(b(a1g1+⋯+akgk−c)p)|]. | (3.6) |
Summing the above formula for t from 1 to k, then the last term of Eq (3.6) is
1pϕk(p−1)2k(p−1)kk∑t=1(kt)T2tpln2tp|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1e(m1g1+n1¯g1+⋯+mtgt+nt¯gtp)⋅e(b(a1g1+⋯+akgk−c)p)|=1pϕk(p−1)2k(p−1)k[kT2pln2p|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1e(m1g1+n1¯g1p)e(b(a1g1+⋯+akgk−c)p)|+⋯+(kk−1)T2(k−1)pln2(k−1)p⋅|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1⋅e(m1g1+n1¯g1+⋯+mk−1gk−1+nk−1¯gk−1p)e(b(a1g1+⋯+akgk−c)p)|+T2kpln2kp|p−1∑b=0p−1∑g1=1⋯p−1∑gk=1e(m1g1+n1¯g1+⋯+mkgk+nk¯gkp)⋅e(b(a1g1+⋯+akgk−c)p)|]≪ϕk(p−1)pk+1ln2kp(pk−12+⋯+pk+12)≪ϕk(p−1)pk+1ln2kp⋅pk−12=ϕk(p−1)p32ln2kp, |
here we have utilized T2p<4π2(1+1.549lnp)2<2.4 and the results in Wang and Wang (see Lemma 2.2 of [13]) that
|p−1∑a=1χd(a)e(ma+n¯ap)|≪p12. |
Similarly, note that ∑d|n|μ(d)|=2ω(n) and we can get the estimate of the other terms of Eq (3.6). Then we have
A2≪ϕk(p−1)p322kω(p−1)ln2kp. | (3.7) |
Inserting Eqs (3.3) and (3.7) into (3.1), we can deduce that
N(R,p)=ϕk(p−1)2kp+O(ϕk(p−1)pk+122kω(p−1))+O(ϕk(p−1)p322kω(p−1)ln2kp)=ϕk(p−1)2kp+O(ϕk(p−1)p322kω(p−1)ln2kp). |
This proves the Theorem 1.
Now we prove the Theorem 2. Let A denote the set of integers 1≤x≤p such that
k∏i=1f(x+ci)≡0(modp). |
By the definition of primitive roots and Lehmer number, it follows that
M(f(x),R,p)=12kϕk(p−1)(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip∑x=1x∉Aχdi(f(x+ci))(1−(−1)f(x+ci)+¯f(x+ci)))=12kϕk(p−1)(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip∑x=1x∉Aχdi(f(x+ci)))+12kϕk(p−1)(p−1)kk∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip∑x=1x∉Aχdi(f(x+ci)))k∑t=1(−1)tk∑i1=1⋯k∑it=1i1<⋯<itgi1⋯git=12kϕk(p−1)(p−1)k(B1+B2), | (3.8) |
where gi=(−1)f(x+ci)+¯f(x+ci),i=1,2,…,k.
B1=k∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip∑x=1x∉Aχdi(f(x+ci)))=p∑x=1x∉A1+k∏i=1(∑di∣p−1k∏i=1di>1μ(di)ϕ(di)∑χdip∑x=1x∉Aχdi(f(x+ci))). |
Obviously,
|p∑x=1x∉A1−p|≤kl. |
From Lemma 6 we have
∑χd1∑χd2⋯∑χdkp∑x=1x∉Aχd1(f(x+c1))χd2(f(x+c2))⋯χdk(f(x+ck))=d1∑s1=1 ′⋯dk∑sk=1 ′p∑x=1x∉Aχ((f(x+c1))s1(p−1)d1⋯(f(x+ck))sk(p−1)dk). |
Due to d1d2⋯dk>1, and
si(p−1)di<p−1 for di>1(i=1,2,…,k), |
from Lemma 4 we can get that the polynomial
(f(x+c1))s1(p−1)d1⋯(f(x+ck))sk(p−1)dk |
has a root in ˉFp with multiples less than p−1, thus it can not be multiple of a (p−1)-th power of polynomial over Fp. Take g(x)=(f(x+c1))s1(p−1)d1⋯(f(x+ck))sk(p−1)dk, in Lemma 2 we have
|p∑x=1x∉Aχ(f(x+c1)s1(p−1)d1⋯f(x+ck)sk(p−1)dk)|<(kl−1)p12. |
Hence, we have
|B1−(p−kl)|<(2kω(p−1)−1)(kl−1)p12≤2kω(p−1)(kl−1)p12. | (3.9) |
Using the methods in the proof of Theorem 1 we have
gi=1p2p−1∑mi,ni=1e(mi(f(x+ci))+ni¯f(x+ci)p)Umi(ui)Uni(vi). |
From the above discussion and Lemma 5, we can obtain
|B2|<|k∏i=1(∑di∣p−1μ(di)ϕ(di)∑χdip∑x=1x∉Aχdi(f(x+ci)))k∑t=1(−1)tk∑i1=1k∑i2=1⋯k∑it=1i1<i2<⋯<itgi1gi2⋯git|<k∏i=1(∑di∣p−1|μ(di)|ϕ(di)∑χdi)k∑t=1(kt)T2tpln2tp⋅|p∑x=1x∉Aχdi(f(x+ci))⋅e(m1(f(x+c1))+n1¯(f(x+c1))+⋯+mt(f(x+ct))+nt¯(f(x+ct))p)|<2kω(p−1)⋅k∑t=1(kt)T2tpln2tp(kl+l)p12. | (3.10) |
Combing Eqs (3.8), (3.9) and (3.10) we have
|M(f(x),R,p)−12kϕk(p−1)(p−1)k(p−kl)|<12kϕk(p−1)(p−1)k[2kω(p−1)(kl−1)p12+2kω(p−1)⋅k∑t=1(kt)T2tpln2tp(kl+l)p12]=12kϕk(p−1)(p−1)k2kω(p−1)p12⋅[(kl−1)+((k+1)l)k∑t=1(kt)T2tpln2tp]. | (3.11) |
Then we have
M(f(x),R,p)=12kϕk(p−1)(p−1)k−1+O(ϕk(p−1)pk−122kω(p−1)ln2kp). |
This complete the proof of Theorem 2.
From two perspectives, this paper consider the distribution of LPRs that are related to the generalized Golomb's conjecture. Theorem 1 extends the binary linear equation ag1+bg2=c to the multivariate linear equation a1g1+a2g2+⋯+akgk=c, and uses the properties of Gauss sums to derive an asymptotic formula for the number of its solutions g1,g2,…,gk that are LPRs. Theorem 2 considers k consecutive LPRs and employs the upper bound estimation of the generalized Kloosterman sums to provide a more general result that for f(x)∈Fp[x], k polynomials f(x+c1),f(x+c2),…,f(x+ck) are Lehmer primitive roots modulo p.
The authors declare they have not used Artificial Intelligence (AI) tools in the creation of this article.
The author gratefully appreciates the referees and academic editor for their helpful and detailed comments.
This work is supported by the N. S. F. (12126357) of P. R. China and the Natural Science Basic Research Plan in Shaanxi Province of China (2023-JC-QN-0050).
The author declare there are no conflicts of interest.
[1] |
O. V. Popovych, C. Hauptmann, P. A. Tass, Control of neuronal synchrony by nonlinear delayed feedback, Biol. Cybern., 95 (2006), 69–85. doi: 10.1007/s00422-006-0066-8
![]() |
[2] | S. Strogatz, Sync, Hyperion, New York, 1 edition, 2003. |
[3] | P. Arkady, Synchronization, Boris Chirikov, Predrag Cvitanovic, Frank Moss, Harry Swinney, New York, 1 edition, 2001. |
[4] |
L. A. Rohde, C. P. Heisenberg, Zebrafish gastrulation: Cell movements, signals, and mechanisms, Int. Rev. Cytol., 261 (2007), 159–192. doi: 10.1016/S0074-7696(07)61004-3
![]() |
[5] | S. Gilbert, Developmental biology, Sinauer Associates, Inc., Publishers, Sunderland, Massachusetts, 2016. |
[6] | T. W. Sadler, Langman's medical embryology, Wolters Kluwer Health/Lippincott Williams & Wilkins, Philadelphia, 2012. |
[7] | B. Carlson, Human embryology and developmental biology, Elsevier/Saunders, Philadelphia, Pa, 2014. |
[8] | Y. Harima, R. Kageyama, Oscillatory links of fgf signaling and hes7 in the segmentation clock. Curr. Opin. Genet. Dev., 23 (2013), 484–490. |
[9] | P. F. Giampietro, S. L. Dunwoodie, K. Kusumi, O. Pourquié, O. Tassy, A. C. Offiah, et al. Progress in the understanding of the genetic etiology of vertebral segmentation disorders in humans, Ann. NY Acad. Sci., 1151 (2008), 38–67. |
[10] |
M. L. Dequéant, O. Pourquié, Segmental patterning of the vertebrate embryonic axis, Nat. Rev. Genet., 9 (2008), 370–382. doi: 10.1038/nrg2320
![]() |
[11] | M. Maroto, R. A. Bone, J. K. Dale, Somitogenesis, Development, 139 (2012), 2453–2456. |
[12] |
J. Cooke, E. C. Zeeman, A clock and wavefront model for control of the number of repeated structures during animal morphogenesis, J. Theor. Biol., 58 (1976), 455–476. doi: 10.1016/S0022-5193(76)80131-2
![]() |
[13] |
I. Palmeirim, D. Henrique, D. Ish-Horowicz, O. Pourquié, Avian hairy gene expression identifies a molecular clock linked to vertebrate segmentation and somitogenesis, Cell, 91 (1997), 639–648. doi: 10.1016/S0092-8674(00)80451-1
![]() |
[14] |
M.-L. Dequeant, E. Glynn, K. Gaudenz, M. Wahl, J. Chen, A. Mushegian, et al. A complex oscillating network of signaling genes underlies the mouse segmentation clock, Science, 314 (2006), 1595–1598. doi: 10.1126/science.1133141
![]() |
[15] |
C. Gomez, E. M. Özbudak, J. Wunderlich, D. Baumann, J. Lewis, O. Pourquié, Control of segment number in vertebrate embryos, Nature, 454 (2008), 335–339. doi: 10.1038/nature07020
![]() |
[16] | H. Y. Wang, Y. X. Huang, L. H. Zheng, Y. L. Bao, L. G. Sun, Y. Wu, et al. Modelling coupled oscillations in the notch, wnt, and FGF signaling pathways during somitogenesis: A comprehensive mathematical model, Comput. Intel. Neurosc., 2015 (2015), 1–16. |
[17] |
J. Lewis, Autoinhibition with transcriptional delay, Curr. Biol., 13 (2003), 1398–1408. doi: 10.1016/S0960-9822(03)00534-7
![]() |
[18] | A. B. Webb, I. M. Lengyel, D. J. Jörg, G. Valentin, F. Jülicher, L. G. Morelli, et al. Persistence, period and precision of autonomous cellular oscillators from the zebrafish segmentation clock, eLife, 5 (2016), e08438. |
[19] | E. M. Özbudak, J. Lewis, Notch signalling synchronizes the zebrafish segmentation clock but is not needed to create somite boundaries, PLoS Genet., 4 (2008), e15. |
[20] |
Y.-J. Jiang, B. L. Aerne, L. Smithers, C. Haddon, D. Ish-Horowicz, J. Lewis, Notch signalling and the synchronization of the somite segmentation clock, Nature, 408 (2000), 475–479. doi: 10.1038/35044091
![]() |
[21] |
J. Lewis, A. Hanisch, M. Holder, Notch signaling, the segmentation clock, and the patterning of vertebrate somites, J. Biol., 8 (2009), 1–7. doi: 10.1186/jbiol111
![]() |
[22] |
W. R. Gordon, K. L. Arnett, S. C. Blacklow, The molecular logic of notch signaling - a structural and biochemical perspective, J. Cell Sci., 121 (2008), 3109–3119. doi: 10.1242/jcs.035683
![]() |
[23] | R. Kopan, Current topics in development biology: Notch Signaling, Academic, San Diego, CA, 2010. |
[24] | E. R. Andersson, U. Lendahl, Therapeutic modulation of notch signalling — are we there yet? Nat. Rev. Drug Discov., 13 (2014), 357–378. |
[25] | S. J. Bray, Notch signalling: a simple pathway becomes complex, Nat. Rev. Mol. Cell Bio., 7 (2006), 678–689. |
[26] |
H. B. Tiedemann, E. Schneltzer, S. Zeiser, I. Rubio-Aliaga, W. Wurst, J. Beckers, et al. Cell-based simulation of dynamic expression patterns in the presomitic mesoderm, J. Theor. Biol., 248 (2007), 120–129. doi: 10.1016/j.jtbi.2007.05.014
![]() |
[27] |
I. H. Riedel-Kruse, C. Muller, A. C. Oates, Synchrony dynamics during initiation, failure, and rescue of the segmentation clock, Science, 317 (2007), 1911–1915. doi: 10.1126/science.1142538
![]() |
[28] |
K. Horikawa, K. Ishimatsu, E. Yoshimoto, S. Kondo, H. Takeda, Noise-resistant and synchronized oscillation of the segmentation clock, Nature, 441 (2006), 719–723. doi: 10.1038/nature04861
![]() |
[29] |
K. Uriu, Y. Morishita, Y. Iwasa, Random cell movement promotes synchronization of the segmentation clock, P.Nat. Acad. Sci., 107 (2010), 4979–4984. doi: 10.1073/pnas.0907122107
![]() |
[30] | O. Cinquin, Repressor dimerization in the zebrafish somitogenesis clock, PLoS Comput. Biol., 3 (2007), e32. |
[31] | C. A. Henry, M. K. Urban, K. K. Dill, J. P. Merlie, M. F. Page, C. B. Kimmel, et al. Two linked hairy/Enhancer of split-related zebrafish genes, her1 and her7, function together to refine alternating somite boundaries, Development, 129 (2002), 3693–3704. |
[32] | A. C. Oates, R. K. Ho, Hairy/E(spl)-related (Her) genes are central components of the segmentation oscillator and display redundancy with the Delta/Notch signaling pathway in the formation of anterior segmental boundaries in the zebrafish, Development, 129 (2002), 2929–2946. |
[33] | C. Schröter, S. Ares, L. G. Morelli, A. Isakova, K. Hens, D. Soroldoni, et al. Topology and dynamics of the zebrafish segmentation clock core circuit, PLoS Biol., 10 (2012), e1001364. |
[34] |
A. Ay, S. Knierer, A. Sperlea, J. Holland, E. M. Ozbudak, Short-lived her proteins drive robust synchronized oscillations in the zebrafish segmentation clock, Development, 140 (2013), 3244–3253. doi: 10.1242/dev.093278
![]() |
[35] |
J.S. Griffith, Mathematics of cellular control processes i. negative feedback to one gene, J. Theor. Biol., 20 (1968), 202–208. doi: 10.1016/0022-5193(68)90189-6
![]() |
[36] |
M. Santillán, On the use of the hill functions in mathematical models of gene regulatory networks, Math. Model. Nat. Pheno., 3 (2008), 85–97. doi: 10.1051/mmnp:2008056
![]() |
[37] |
D. Gonze, S. Bernard, C. Waltermann, A. Kramer, H. Herzel, Spontaneous synchronization of coupled circadian oscillators, Biophys. J., 89 (2005), 120–129. doi: 10.1529/biophysj.104.058388
![]() |
[38] |
J. Garcia-Ojalvo, M. B. Elowitz, S. H. Strogatz, Modeling a synthetic multicellular clock: Repressilators coupled by quorum sensing, P. Nat. Acad. Sci., 101 (2004), 10955–10960. doi: 10.1073/pnas.0307095101
![]() |
[39] | W. Wu, W. J. Zhou, T. P. Chen, Cluster synchronization of linearly coupled complex networks under pinning control, IEEE T. Circuits I, 56 (2009), 829–839. |
[40] |
T. Dahms, J. Lehnert, E. Schöll, Cluster and group synchronization in delay-coupled networks, Phys. Rev. E, 86 (2012), 016202. doi: 10.1103/PhysRevE.86.016202
![]() |
[41] |
F. Sorrentino, L. Pecora, Approximate cluster synchronization in networks with symmetries and parameter mismatches, Chaos: An Interdisciplinary Journal of Nonlinear Science, 26 (2016), 094823. doi: 10.1063/1.4961967
![]() |
[42] |
J. W. Feng, P. Yang, Y. Zhao, Cluster synchronization for nonlinearly time-varying delayed coupling complex networks with stochastic perturbation via periodically intermittent pinning control, Appl. Math. Comput., 291 (2016), 52–68. doi: 10.1016/j.amc.2016.06.030
![]() |
[43] |
C. Ma, Q. R. Yang, X. Q. Wu, J. A. Lu, Cluster synchronization: From single-layer to multi-layer networks, Chaos: An Interdisciplinary Journal of Nonlinear Science, 29 (2019), 123120. doi: 10.1063/1.5122699
![]() |
[44] |
G. B. Ermentrout, Oscillator death in populations of "all to all" coupled nonlinear oscillators, Physica D, 41 (1990), 219–231. doi: 10.1016/0167-2789(90)90124-8
![]() |
[45] |
F. A. Rodrigues, T. K. DM. Peron, P. Ji, J. Kurths, The kuramoto model in complex networks, Physics Reports, 610 (2016), 1–98. doi: 10.1016/j.physrep.2015.10.008
![]() |
[46] |
J. Gómez-Gardeñes, Y. Moreno, A. Arenas, Synchronizability determined by coupling strengths and topology on complex networks, Phys. Rev. E, 75 (2007), 066106. doi: 10.1103/PhysRevE.75.066106
![]() |
[47] |
E. A. Delaune, P. François, N. P. Shih, S. L. Amacher, Single-cell-resolution imaging of the impact of notch signaling and mitosis on segmentation clock dynamics, Dev. Cell, 23 (2012), 995–1005. doi: 10.1016/j.devcel.2012.09.009
![]() |
[48] |
J.-N. Teramae, H. Nakao, G. B. Ermentrout, Stochastic phase reduction for a general class of noisy limit cycle oscillators, Phys. Rev. Lett., 102 (2009), 194102. doi: 10.1103/PhysRevLett.102.194102
![]() |
[49] |
K. Kotani, I. Yamaguchi, Y. Ogawa, Y. Jimbo, H. Nakao, G. B. Ermentrout, Adjoint method provides phase response functions for delay-induced oscillations, Phys. Rev. Lett., 109 (2012), 044101. doi: 10.1103/PhysRevLett.109.044101
![]() |