Abstract
Phyllotactic patterns in plants are well known to be related to the golden ratio. Actually, many mathematical models using the theoretical inhibitory effect were proposed to reproduce these phyllotactic patterns. In 1996, Douady and Couder introduced a model using magnetic repulsion and succeeded in reproducing phyllotactic patterns numerically. On the other hand, it was recently revealed in biological experiments that a plant hormone, auxin, regulates the phyllotactic formation as an activator (Reinhardt et al., Nature 426:255–260, 2003). Then, there arises a natural question as to how the inhibitory effect can be related to the auxin. In this paper, a reaction diffusion model is proposed by taking account of auxin behavior in plant tips. The relationship between Douady and Couder’s model and our model is shown by singular limit analysis. It also provides us with the potential function corresponding to the inhibitory effect, and the bifurcation diagram.
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Acknowledgments
The authors would like to thank Professor Masahiko Furutani of Nara Institute of Science and Technology for discussions about the auxin transport and biological experiments, and Professor Ryo Kobayashi of Hiroshima University for the fruitful discussions. The authors are particularly grateful to the referees for their valuable comments. YT has been supported by Meiji University MIMS Ph.D. Program. MM is partially supported by JSPS KAKENHI Grant Nos. 15K13462 and HN is partially supported by JSPS KAKENHI Grant Nos. 25610036 and 26287024.
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Appendix
Appendix
Here we will provide an outline of the proof of the theorem and explain the key proposition to overcome the difficulty in the proof. We divide \(S_n^{\varepsilon }(x,t)\) defined in (6) into the following two parts:
To show that \(S^{ \varepsilon }_n(x,t) \) is the weak solution of \((M_n^{\varepsilon })\), we introduce the following auxiliary system:
where \(j=0, \ldots , n-1\), \(n \ge 1\), and \(\rho _{\eta }(x)\) is the Friedrichs mollifier with a small parameter \(0<\eta \ll 1\) defined by
The initial conditions for \((S_n, \{X_j^{\varepsilon ,\eta } \}_{j=0}^{2})\), are set similarly to those of \((M_n^{\varepsilon })\). Using the heat kernel, we have the classical solution of \((M_n^{\varepsilon ,\eta })\),
Theorem A.1
Assume that
- \((a1)_n\) :
-
\(S^{\varepsilon , \eta }_{n-1}(\cdot , (n-1)T), \ S^{\varepsilon }_{n-1}(\cdot , (n-1)T) \in L^{1}({{\mathbb {R}}}^2)\),
- \((a2)_n\) :
-
\(\sup _{\varepsilon ,\eta >0} \Vert S^{\varepsilon ,\eta }_{n-1}(\cdot , (n-1)T) \Vert _{L^1({{\mathbb {R}}}^2)} <C_1\),
- \((a3)_n\) :
-
\( \sup _{\varepsilon >0} \Vert S^{\varepsilon }_{n-1}(\cdot , (n-1)T) \Vert _{L^1({{\mathbb {R}}}^2)} <C_2\),
- \((a4)_n\) :
-
\(\left\| S^{\varepsilon , \eta }_{n-1}(\cdot , (n-1)T) - S^{\varepsilon }_{n-1}(\cdot , (n-1)T) \right\| _{L^1({{\mathbb {R}}}^2)} \rightarrow 0 \ as \ \eta \rightarrow +0\),
- \((a5)_n\) :
-
\(\theta _{j }^{\varepsilon , \eta } \rightarrow \theta _{j }^{\varepsilon } \ as \ \eta \rightarrow +0 \ (j=2, \ldots , n-1)\),
- \((a6)_n\) :
-
\(\theta _{j }^{\varepsilon } \rightarrow \theta _{j }^{0} \ as \ \varepsilon \rightarrow +0 \ (j=2, \ldots , n-1)\).
Then \(S_n^{\varepsilon }(x,t)\) is the unique weak solution of \((M_n^{\varepsilon })\) for \(t \in ((n-1)T, nT]\). Moreover,
for \(t \in ((n-1)T, nT]\) and
as \(\varepsilon \rightarrow +0\). Additionally, if \(\theta \in [0,2\pi )\) attaining the minimum of \({ \tilde{S}}^{0}_n(\theta )\) on \(C_{R_0}(\theta )\) is unique,
as \(\varepsilon \rightarrow +0\).
Proof of Theorem 4.1
Theorem 4.1 follows from Theorem A.1 via the induction. Indeed, \(S_1^{\varepsilon }(x,t)\) and \(S_1^{0}(x,t)\) satisfy the all assumptions \((a1)_1\)–\((a6)_1\) for (0, T]. Hence, we obtain that \(S_1^{\varepsilon }(x,t)\) is the unique weak solution of \((M_1^{\varepsilon })\), \(\left\| S^{\varepsilon }_1(\cdot , t) - g_1(\cdot , t) \right\| _{L^1({\mathbb {R}}^2)} \rightarrow 0 \) for (0, T] as \(\varepsilon \rightarrow +0\), \(\left\| { \tilde{S}}_{1}^{\varepsilon }( \cdot ) - { \tilde{g}_1}(\cdot ) \right\| _{C^1( [0, 2\pi ) )} \rightarrow 0 \) and \(\theta ^{\varepsilon }_1 \rightarrow \theta ^{0}_1\) as \(\varepsilon \rightarrow +0\) since \(\theta _1^0\) is unique at \(t=T\). Thus, the first step of the induction is established. Next suppose that the theorem for \(( 0, (n-1)T]\) holds. All assumptions \((a1)_n\)–\((a6)_n\) can be confirmed by Theorem A.1. Therefore, by induction, the proof of Theorem 4.1 is completed. \(\square \)
Proof of Theorem A.1
We mainly explain the outline of the proof. From the assumptions \((a1)_n\)–\((a5)_n\), we can prove \(\Vert S_n^{\varepsilon ,\eta }(\cdot , t)- S_n^{\varepsilon }(\cdot , t) \Vert _{L^1({{\mathbb {R}}}^2)} \rightarrow 0\) as \(\eta \rightarrow +0\) by the continuity of \(\varPsi _h\) in \(L^1({{\mathbb {R}}}^2)\) where \(\varPsi _h\) is the shift operator defined by \(\varPsi _hf(\cdot ):=f(x-h)\) for \(x,h \in {{\mathbb {R}}}^2\), namely, \(\Vert \varPsi _h f(\cdot ) -f(\cdot ) \Vert _{L^1({{\mathbb {R}}}^2)} \rightarrow 0\) as \(h \rightarrow 0\). It also follows from \((a1)_n\), \((a4)_n\), \((a5)_n\) and the dominated convergence theorem that \(\Vert {\tilde{S}}_n^{\varepsilon ,\eta }(\cdot )- {\tilde{S}}_n^{\varepsilon }(\cdot ) \Vert _{C^1([0, 2\pi ))} \rightarrow 0\) as \(\eta \rightarrow +0\). Thus, we can determine that \(\theta _n^{\varepsilon , \eta } \rightarrow \theta ^{\varepsilon }\) as \(\eta \rightarrow +0\). Using \(\Vert S_n^{\varepsilon ,\eta }(\cdot , t)- S_n^{\varepsilon }(\cdot , t) \Vert _{L^1({{\mathbb {R}}}^2)} \rightarrow 0\) for \(t \in [(n-1)T, nT]\) and \((a5)_n\), we can show that \(S_n^{\varepsilon }(x, t)\) is the unique weak solution of \((M_n^{\varepsilon })\) for \(t \in ((n-1)T, nT]\) by the Hölder inequality. From \((a3)_n\) and \((a6)_n\) we can show \(\Vert {\tilde{S}}_n^{\varepsilon }(\cdot )- {\tilde{S}}_n^{0}(\cdot ) \Vert _{C^1([0, 2\pi ))} \rightarrow 0\) as \(\varepsilon \rightarrow +0\) which implies (A.1). Since \(S_n^{\varepsilon }\) has the singularity points moving along \(X^{\varepsilon }_j( t - \varepsilon \tau ) (j=0,\ldots ,n-1)\) for \(\tau \in [0, (t-(n-1)T)/\varepsilon ]\), we cannot apply the dominated convergence theorem to show (7). We prepare the following proposition. \(\square \)
Proposition A.1
Under the assumptions \((a3)_n\) and \((a6)_n\),
as \(\varepsilon \rightarrow +0\) for \(t \in ((n-1)T,nT]\).
Proof
It is easily shown that \(S_n^{\varepsilon }( \cdot ,t),S_n^{0}( \cdot ,t) \in L^1({\mathbb {R}}^2)\). Therefore, we omit it. Next, we show \(\Vert S^{\varepsilon }_n(\cdot ,t) - S^{0}_n(\cdot ,t) \Vert _{L^1({\mathbb {R}}^2)} \rightarrow 0\) as \(\varepsilon \rightarrow +0\). Note that \(S_n^{\varepsilon }( x,t) - S_n^{0}( x,t) = P_n^{\varepsilon }( x,t) + N_n^{\varepsilon }( x,t) - S_n^{0}( x,t) \). Since
as \(\varepsilon \rightarrow +0\), we only need to estimate \(\Vert N_n^{\varepsilon }( \cdot ,t) - S^{0}_n( \cdot ,t) \Vert _{L^1({\mathbb {R}}^2)}\). We have that
We denote the first and second terms on the right hand side by \(Q_n^{\varepsilon }(x,t)\) and \(R_n^{\varepsilon }(x,t)\), respectively. We calculate that
as \(\varepsilon \rightarrow +0\). Because \(N^{\varepsilon }_n(x,t)\) and \(S^{0}_n(x,t)\) have singularities at \(X_j^{\varepsilon }(t-\varepsilon \tau )\) and \(X_j^{0}(t)\), respectively, we divide \(\Vert Q_n^{\varepsilon } \Vert _{L^1({{\mathbb {R}}}^2)}\) into three parts.
where \(0<\sigma \ll 1\) and
Firstly, we estimate \( H_n^{\varepsilon }( t)\). As \(C_0( {{\mathbb {R}}}^2 \times [0, \sigma ])\) is dense in \(L^1( {{\mathbb {R}}}^2 \times [0, \sigma ])\), for any \({\tilde{\varepsilon }} > 0 \) there exists a function \(\xi (\cdot ,\cdot ) \in C_0( {{\mathbb {R}}}^2 \times [0, \sigma ])\) such that \(\Vert K(\cdot ,\cdot ) - \xi (\cdot ,\cdot ) \Vert _{L^1({{\mathbb {R}}}^2 \times [0,\sigma ))} < {\tilde{\varepsilon }}\). Moreover, for any \({ \tilde{\varepsilon }}>0\), there exists \({ \tilde{\delta }}>0\), such that whenever \( \left| X^0_j(t )-X^{\varepsilon }_j(t -\varepsilon \tau ) \right| < { \tilde{\delta }} \) for \(\tau \in [0, \sigma ]\), we have \(\left| \varPsi _{X^0_j(t )-X^{\varepsilon }_j(t -\varepsilon \tau ) }\xi (\cdot , \tau ) - \xi (x, \tau ) \right| < { \tilde{\varepsilon }} \) since \(\theta ^{\varepsilon } _j \rightarrow \theta ^{0}_j\) as \(\varepsilon \rightarrow +0 \ (j=1, \ldots , n-1)\). Thus we have
Secondly, we consider \(I^{\varepsilon }_n(t)\). If \(x \in \varOmega _{\varepsilon }\),
where
Thus, using \((a6)_n\), we compute
as \(\varepsilon \rightarrow 0\). Finally,we consider \(J_n^{\varepsilon }(t)\). Set
Then
where \(\chi _{\varTheta _n}(x,\tau )\) is a characteristic function and
As a result of the characteristic function \(\chi _{\varTheta _n}(x,\tau )\), the domain of integration \(J_n^{\varepsilon }(t)\) is independent of \(\varepsilon \). Thus, we have that \(f_n^{\varepsilon }(x,\tau )\chi _{\varTheta _n}(x,\tau ) \rightarrow 0\) pointwise in \( {{\mathbb {R}}}^2 \times [\sigma , \infty ]\) as \(\varepsilon \rightarrow +0\). Since \(\left| x - X^{\varepsilon }_j(t - \varepsilon \tau ) \right| > \left| x - X^0_j(t) \right| /2\) for \(x \notin \varOmega _{\varepsilon }, \ 0 \le \tau < (t-(n-1)T)/\varepsilon \), we have
which is measurable in \({{\mathbb {R}}}^2 \times [\sigma ,\infty )\). Thus from the dominated convergence theorem, we compute that
\(\square \)
This proposition immediately implies (7). Thus, we have completed the proof of Theorem A.1. \(\square \)
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Tanaka, Y., Mimura, M. & Ninomiya, H. A reaction diffusion model for understanding phyllotactic formation. Japan J. Indust. Appl. Math. 33, 183–205 (2016). https://doi.org/10.1007/s13160-015-0202-8
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DOI: https://doi.org/10.1007/s13160-015-0202-8