Abstract
This paper is concerned with a nonlinear optimization problem that naturally arises in population biology. We consider the population of a single species with logistic growth residing in a patchy environment and study the effects of dispersal and spatial heterogeneity of patches on the total population at equilibrium. Our objective is to maximize the total population by redistributing the resources among the patches under the constraint that the total amount of resources is limited. It is shown that the global maximizer can be characterized for any number of patches when the diffusion rate is either sufficiently small or large. To show this, we compute the first variation of the total population with respect to resources in the two patches case. In the case of three or more patches, we compute the asymptotic expansion of all patches by using the Taylor expansion with respect to the diffusion rate. To characterize the shape of the global maximizer, we use a recurrence relation to determine all coefficients of all patches.
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YL was partially supported by the NSF grant DMS-1853561; EY was partially supported by JSPS KAKENHI Grant-in-Aid for Scientific Research (A) (No. 17H01095).
Appendix
Appendix
1.1 Preliminaries for small \(\delta \)
In this Appendix, we choose \(N \ge 2\) and \(\varvec{}{m} \in \mathcal {M}\) arbitrarily.
Lemma 7
Any solution of (1.3) satisfies either \({{\,\mathrm{\underline{deg}}\,}}u_i = 0\) or \(1/2 \le {{\,\mathrm{\underline{deg}}\,}}u_i < 1\) for each \(i \in \varOmega \). Moreover, \(m_i > 0\) if \({{\,\mathrm{\underline{deg}}\,}}u_i = 0\), and \(m_i = 0\) if \(1/2 \le {{\,\mathrm{\underline{deg}}\,}}u_i < 1\).
Proof
By using (3.1)–(3.3), \(u_i \rightarrow m_i\) as \(\delta \rightarrow 0\). This implies \({{\,\mathrm{\underline{deg}}\,}}u_i \ge 0\). First, we choose \(i = 1\) or \(i = N\). Assume \(m_i > 0\). Then by (3.1), we have
Similarly, by (3.3), we have
This indicates that if \(m_i > 0\), then \({{\,\mathrm{\underline{deg}}\,}}u_i = 0\).
Next, assume \(m_i = 0\). By using (3.1) and (3.3), we have
This gives
This argument and Proposition 1 indicate that if \(m_i = 0\), then \(1/2 \le {{\,\mathrm{\underline{deg}}\,}}u_i \le 1\). It follows that \({{\,\mathrm{\underline{deg}}\,}}u_i = 0\) is equivalent to \(m_i > 0\). From the above argument, we can assert that \(1/2 \le {{\,\mathrm{\underline{deg}}\,}}u_i \le 1\) is equivalent to \(m_i = 0\).
Second, we choose \(i \in \mathbb {Z} \cap (1,N)\). Assume \(m_i > 0\). By (3.2), we have
This implies that if \(m_i > 0\), then \({{\,\mathrm{\underline{deg}}\,}}u_i = 0\). Next, assume \(m_i = 0\). We use (3.2) again to have
This gives
Similar argument to the first case shows that \({{\,\mathrm{\underline{deg}}\,}}u_i = 0\) is equivalent to \(m_i > 0\). Furthermore, \(1/2 \le {{\,\mathrm{\underline{deg}}\,}}u_i \le 1\) is equivalent to \(m_i = 0\).
Therefore, it is sufficient to prove that \(0 \le {{\,\mathrm{\underline{deg}}\,}}u_i < 1\) for all \(i \in \varOmega \). We assume that there exists \(i \in \varOmega \) such that \({{\,\mathrm{\underline{deg}}\,}}u_i = 1\). From (A.1)–(A.3), we have
Repeated application of (A.1)–(A.3) enables us to obtain \(m_i = 0\) for all \(i \in \varOmega \), which contradicts our assumption. \(\square \)
Recall that (6.1)–(6.3) are obtained by Lemma 7. We next show that \({{\,\mathrm{\underline{deg}}\,}}u_i\) takes a finite number of values. By Lemma 8, we only have to seek the largest coefficients of an appropriate order of \(\delta \) to maximize the total population.
Lemma 8
Any solution of (1.3) satisfies
for all \(i \in \varOmega \).
Proof
We assume that there exists \(i \in \varOmega \) such that
where \(p \in \mathbb {Z} \cap [0, \min \{ i-1, N-i \}]\). By using (A.1)–(A.3), we calculate
Repeating substitution of (A.4) to (A.5) p times, we obtain
This contradicts Lemma 7. Finally, we assume that
Applying (A.1)–(A.3) repeatedly \(\max \{ N-i , i-1 \}\) times to have \(\min _{i \in \varOmega } \{ {{\,\mathrm{\underline{deg}}\,}}u_i\}\). Since \(\max \{ N-i , i-1 \} \le N-1\), we have \(\min _{i \in \varOmega } \{ {{\,\mathrm{\underline{deg}}\,}}u_i\} > 0\). This contradicts our assumption. \(\square \)
We next evaluate the difference of \({{\,\mathrm{\underline{deg}}\,}}u_i\) between two adjacent patches.
Lemma 9
Let \(i \in \varOmega \) and \(p \in \mathbb {Z} \cap [0, N-1]\). Suppose that a solution of (1.3) satisfies
Then we have
where the equality holds if and only if \(m_i = 0\), and
where the equality holds if and only if \(\min \{ m_{i-1}, m_{i+1} \} = 0\).
Proof
It is clear that (A.6) follows immediately from (A.5). Suppose that
By (A.6), we have
This contradicts our assumption. This proves (A.7). \(\square \)
Repeated application of Lemma 9 enables us to have the following lemma.
Lemma 10
Let \(i,k \in \varOmega \), and take \(p \in \mathbb {Z} \cap [1, N-1]\) such that \(k-p\), \(k+p \in \varOmega \). If a positive solution of (1.3) satisfies \({{\,\mathrm{\underline{deg}}\,}}u_k = (2^p - 1)/2^p\), then
Moreover, \(m_i = 0\) for all \(i \in ( k-p, k+p)\).
Lemma 10 allows us to examine the effect of the distance between favorable patches on \({{\,\mathrm{\underline{deg}}\,}}u_i\). The following proposition plays an important role in Sect. 6.
Lemma 11
Let k, i, q be positive integers such that \(k, k+i, k+2q \in \varOmega \).
-
(i)
Suppose that \(m \in \mathcal {M}\) satisfies \(m_k > 0\), \(m_{k+2q} > 0\), and \(m_{k+i} = 0 \; (0<i< 2q)\). Then the following equalities hold:
$$\begin{aligned} {{\,\mathrm{\underline{deg}}\,}}u_{k+i} = {{\,\mathrm{\underline{deg}}\,}}u_{k+2q-i } = \frac{2^i - 1}{2^i} \quad (0 \le i \le q). \end{aligned}$$ -
(ii)
Suppose that \(\varvec{}{m} \in \mathcal {M}\) satisfies \(m_k > 0\), \(m_{k+2q} > 0\), and \(m_{k+i} = 0 \; (0<i< 2q+1)\). Then the following equalities hold:
$$\begin{aligned} {{\,\mathrm{\underline{deg}}\,}}u_{k+i} = {{\,\mathrm{\underline{deg}}\,}}u_{k+2q+1-i} = \frac{2^i - 1}{2^i} \quad (0 \le i \le q). \end{aligned}$$
Proof
(i) Assume that \(i \in \mathbb {Z} \cap [1,q]\) and
where \(j \in \mathbb {Z} \cap [1, i]\). Then we have
by Lemma 10. Similarly, assuming that
where \(j \in \mathbb {Z} \cap [1, N-1-i]\), we have \(m_k = 0\) by Lemma 10. This contradicts our assumption \(m_k>0\). We now apply this argument again, with \(k+i\) replaced by \(k + 2q - i\), to obtain the value of \({{\,\mathrm{\underline{deg}}\,}}u_{k+2q-i}\).
The case (ii) may be proved in much the same way as the case (i), so we omit the proof. \(\square \)
1.2 Expansion of the total population in N patches
To express the total population \(U(\varvec{}{m}, \delta )\) in terms of \(\delta \) for each \(\varvec{}{m} \in \mathcal {M}\), we show the following result.
Proposition 7
Let k, i, q be positive integers such that \(k, k+i, k+2q \in \varOmega \). Choose \(\varvec{}{m} \in \mathcal {M}\) arbitrarily. Then the positive solution of (1.3) are expanded as follows:
- (i):
-
If \(\varvec{}{m} \in \mathcal {M}\) satisfies \(m_k > 0\), \(m_{k+2q} > 0\) and \(m_{k+i} = 0, \; (1 \le i \le 2q-1)\), then
$$\begin{aligned} u_{k}&= {\left\{ \begin{array}{ll} m_k - \delta +o(\delta ), &{}\; (k = 1),\\ \displaystyle m_k + \left( \frac{m_{k-1}}{m_k} - 2 \right) \delta + o(\delta ), &{}\; (k>1), \end{array}\right. } \end{aligned}$$(A.8)$$\begin{aligned} u_{k+i}&= m_k^{1/2^i} \delta ^{(2^i - 1)/2^{-i}} - \delta + o(\delta ), \; (1 \le i \le q-1), \end{aligned}$$(A.9)$$\begin{aligned} u_{k+q}&= \sqrt{m_k^{1/2^{q-1}} + m_{k+2q}^{1/2^{q-1}}} \delta ^{(2^q - 1)/2^q} - \delta + o(\delta ), \end{aligned}$$(A.10)$$\begin{aligned} u_{k+2q-i}&= m_{k+2q}^{1/2^i} \delta ^{(2^i - 1)/2^i} - \delta + o(\delta ), \; (1 \le i \le q-1), \end{aligned}$$(A.11)$$\begin{aligned} u_{k+2q}&= {\left\{ \begin{array}{ll} m_{k+2q} - \delta +o(\delta ), &{}\; (k+2q = N), \\ \displaystyle m_{k+2q} + \left( \frac{m_{k+2q+1}}{m_{k+2q}} - 2 \right) \delta + o(\delta ), &{}\; (k+2q<N). \end{array}\right. } \end{aligned}$$(A.12) - (ii):
-
If \(\varvec{}{m} \in \mathcal {M}\) satisfies \(m_k > 0\), \(m_{k+2q+1} > 0\) and \(m_{k+i} = 0, \; (1 \le i \le 2q)\), then
$$\begin{aligned} u_{k}= & {} {\left\{ \begin{array}{ll} m_k - \delta +o(\delta ), &{}\; (k = 1),\\ \displaystyle m_k + \left( \frac{m_{k-1}}{m_k} - 2 \right) \delta + o(\delta ), &{}\; (k>1), \end{array}\right. }\nonumber \\ u_{k+i}= & {} m_k^{1/2^i} \delta ^{(2^i - 1)/2^i} - \delta + o(\delta ), \; (1 \le i \le q-1),\nonumber \\ u_{k+q}= & {} m_k^{1/2^{q}} \delta ^{(2^q - 1)/2^q} + \left( \frac{1}{2} \left( \frac{m_{k+2q+1}}{m_k} \right) ^{1/2^{q}} -1 \right) \delta + o(\delta ), \end{aligned}$$(A.13)$$\begin{aligned} u_{k+q+1}= & {} m_{k+2q+1}^{1/2^{q}} \delta ^{(2^q - 1)/2^q} + \left( \frac{1}{2} \left( \frac{m_k}{m_{k+2q+1}} \right) ^{1/2^{q}} -1 \right) \delta + o(\delta ), \nonumber \\ u_{k+2q+1-i}= & {} m_{k+2q+1}^{1/2^i} \delta ^{(2^i - 1)/2^i} - \delta + o(\delta ), \; (1 \le i \le q-1),\nonumber \\ u_{k+2q+1}= & {} {\left\{ \begin{array}{ll} m_{k+2q+1} - \delta +o(\delta ), &{}\; (k+2q+1 = N), \\ \displaystyle m_{k+2q+1} + \left( \frac{m_{k+2q+2}}{m_{k+2q+1}} - 2 \right) \delta + o(\delta ), &{}\; (k+2q+1<N). \end{array}\right. } \end{aligned}$$(A.14) - (iii):
-
If \(\varvec{}{m} \in \mathcal {M}\) satisfies \(m_k> 0, \; (k > 1)\), \(m_i = 0, \; (1 \le i \le k-1)\), then
$$\begin{aligned} u_{1}= & {} m_k^{1/2^{k-1}} \delta ^{(2^{k-1} - 1)/2^{k-1}} - \frac{\delta }{2} + o(\delta ),\nonumber \\ u_{k-i}= & {} m_k^{1/2^{i}} \delta ^{(2^{i} - 1)/2^{i}} - \delta + o(\delta ),\nonumber \\ u_{k}= & {} {\left\{ \begin{array}{ll} m_k - \delta +o(\delta ), &{}\; (k = N),\\ \displaystyle m_k + \left( \frac{m_{k+1}}{m_k} - 2 \right) \delta + o(\delta ), &{}\; (k<N), \end{array}\right. } \end{aligned}$$(A.15) - (iv):
-
If \(\varvec{}{m} \in \mathcal {M}\) satisfies \(m_k > 0, \; (k < N)\), \(m_i = 0, \; (k +1 \le i \le N)\), then
$$\begin{aligned} \begin{aligned} u_{k}&= {\left\{ \begin{array}{ll} m_k - \delta +o(\delta ), &{}\; (k = 1),\\ \displaystyle m_k + \left( \frac{m_{k-1}}{m_k} - 2 \right) \delta + o(\delta ), &{}\; (k>1), \end{array}\right. }\\ u_{k+i}&= m_k^{1/2^{i}} \delta ^{(2^{i} - 1)/2^{i}} - \delta + o(\delta ) ,\\ u_{N}&= m_k^{1/2^{N-k}} \delta ^{(2^{N-k} - 1)/2^{N-k}} - \frac{\delta }{2} + o(\delta ). \end{aligned} \end{aligned}$$
Proof
(i) It is easy to verify (A.8) by noting \({{\,\mathrm{\underline{deg}}\,}}u_{k+1} = 1/2\). So we omit the proof. We now consider (A.9) by induction on i. By Lemma 11, there exist coefficients \(C_{k+i}\) for all \(i \in \{0, 1, \ldots , q\}\) such that
By (6.1) or (6.2), we have the base case as
Therefore, we have the relation between \(C_{k}\) and \(C_{k+1}\) given by
Suppose that
By (6.2), we have the induction step as
Therefore, we have recurrence relation \(C_{k+i} = C_{k+i-1}^{1/2}\) which gives (A.9). From this, we replace “k” by “\(k+2q\)” to have (A.11) and (A.12).
To prove (A.10), we use (A.9) and (A.11) to have
By (6.2), we obtain
(ii) We give the proof only for (A.13) and (A.14); the other cases can be proved by the same argument as above. We use Lemma 11 to have
We now compute \(C_{k+q}\) and \(C_{k+q+1}\) by using (6.2) as
Similarly, we have
By the definition of \(C_{k+q}\) and \(C_{k+q+1}\), we have
Therefore, we obtain (A.13) and (A.14).
(iii) We use (6.1) to have
This proves (A.15) The other cases can be proved by the same argument.
(iv) This can be proved in much the same way as (iii). So we omit the proof. \(\square \)
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Nagahara, K., Lou, Y. & Yanagida, E. Maximizing the total population with logistic growth in a patchy environment. J. Math. Biol. 82, 2 (2021). https://doi.org/10.1007/s00285-021-01565-7
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DOI: https://doi.org/10.1007/s00285-021-01565-7