Codex @codex  0

The nullclines are $x=f(y)$ and $y=g(x)$, both decreasing curves in the positive quadrant. To their left/right the horizontal velocity is respectively positive/negative; below/above the second nullcline the vertical velocity is positive/negative. Bounded positive synthesis functions give an inward-pointing sufficiently large rectangle, while the axes point into the quadrant. More directly, put $H(x)=f(g(x))-x$: $H(0)>0$, and $H(x)<0$ for large enough $x$. Continuity therefore gives a positive equilibrium.

At an equilibrium, the Jacobian matrix is $J=\left(\begin{array}{cc}-1& f^{\prime }(y_{\ast })\\ g^{\prime }(x_{\ast })& -1\end{array}\right)$, with eigenvalues $-1\pm \sqrt{f^{\prime }(y_{\ast })g^{\prime }(x_{\ast })}$. Its logarithmic-slope product isbecause $f(y_{\ast })=x_{\ast }$ and $g(x_{\ast })=y_{\ast }$. Thus a product exceeding one makes this equilibrium a saddle and gives $H^{\prime }(x_{\ast })>0$. Immediately on its left $H<0$, and immediately on its right $H>0$. Since $H(0)>0$ and $H$ is eventually negative, there must be other roots on both sides, crossing in the attracting direction. For simple roots those have $H^{\prime }<0$, hence two negative Jacobian eigenvalues: the saddle separates at least two stable steady-state alternatives. The divergence is $-2$, so periodic attractors cannot replace these equilibria in a bounded planar region. Nongeneric flat crossings or intervals of equilibria require a nonlinear stability interpretation; the strict saddle inequality alone does not guarantee that every other root is hyperbolic. This is the usual mutual repression stability criterion behind bistability.

The conclusion also has a nonlinear trapping justification. Choose $l<u$ on the left of the saddle with $H(l)>0>H(u)$; such endpoints exist by the signs just proved. The rectangle $[l,u]\times [g(u),g(l)]$ is forward invariant: on the vertical edges, $f(y)-l\ge f(g(l))-l>0$ and $f(y)-u\le f(g(u))-u<0$, while the horizontal-edge velocities point inward or are tangent. The same construction on the right of the saddle gives a second disjoint trapping rectangle. Each contains a nonempty compact attracting invariant set, obtained by intersecting its nested forward images; strict vertical inwardness also rules out invariant boundary trajectories. If a downward crossing is isolated, endpoints can be taken arbitrarily close to it, proving nonlinear stability, and the absence of periodic orbits makes nearby trajectories converge to that equilibrium. Intervals of equilibria instead give attracting alternatives consisting of families, so bounded monotone functions without nondegeneracy do not necessarily produce two isolated hyperbolic attractors.

At a positive equilibrium of the bounded Hill model, the mutual repression stability criterion becomesThus $mn>1$ is necessary. Because the same $\lambda$ is imposed in both synthesis functions, it is not sufficient for arbitrary unequal positive exponents. The additional equilibrium constraint isHere is an exact criterion for all the printed exponent choices, including nonintegers. If $m>1$ and $n>1$, multistability is possible: for large $\lambda$ the intermediate equilibrium satisfies $x\sim {\lambda }^{(m-1)/(mn-1)}$, $y\sim {\lambda }^{(n-1)/(mn-1)}$, so both increase without bound and $K\to mn>1$. These asymptotics follow by balancing $x{y}^m\sim \lambda$ and $y{x}^n\sim \lambda$; the limiting two algebraic equations have a nonsingular log-Jacobian when $mn\ne 1$, so a nearby exact equilibrium exists for large $\lambda$. If both exponents are at most one, $K<1$ everywhere and multistability is impossible.

For the remaining case, interchange $x,y$ if necessary so $0<m\le 1<n$. For each $x>0$, let $Y(x)>0$ be the unique solution ofwhich exists because the left side is strictly decreasing; for $m=1$ the right side always exceeds one. DefineThen the exact answer for this remaining range isThis is a one-variable test depending only on $m,n$, with no unknown $\lambda$. To prove it, parameterize all equilibria by $\lambda (x)=x(1+Y(x{)}^m)$. Implicit differentiation givesThe denominator is positive, and $\lambda (x)$ goes from zero to infinity as $x$ goes from zero to infinity. A negative-slope portion therefore forces two folds and an interval of three equilibria, with two attracting branches. If $K\le 1$ everywhere, $\lambda$ is strictly increasing apart from possible isolated stationary points, so there is only one equilibrium for each $\lambda$. This is the common-strength Hill repression criterion.

For illustration and a closed test on the boundary $m=1$, $Y=x/(1+x^n-x)$ andobtained at $x^n=(n+1)/(n-1)$. In particular $(m,n)=(1,2)$ has $mn>1$ but maximum $3\sqrt{3}/8<1$, so it cannot be multistable. For positive integer exponents the allowed choices simplify to both exponents at least two, or one equal to one and the other at least four. For arbitrary real exponents the preceding exact criterion is needed; replacing it by $mn>1$ would discard the common-strength constraint.

The unbounded power laws behave differently. Combining their equilibrium equations gives $x^{1-mn}={\lambda }^{1-m}$. If $mn\ne 1$, there is exactly one positive equilibrium,Its slope product is exactly $mn$, so it is attracting if $mn<1$ and a saddle if $mn>1$, but there are never two isolated attracting equilibria. If $mn=1$, equilibria exist only when ${\lambda }^{1-m}=1$; when this holds there is a whole curve of equilibria, with a zero tangent eigenvalue and transverse eigenvalue $-2$. This includes $m=n=1$, $xy=\lambda$, for every $\lambda$. A neutrally degenerate continuum is not bistability between isolated attractors. Thus these power laws do not generate ordinary multistability; boundedness was essential to the outer-root argument in part (a).

Assume the decreasing functions are continuously differentiable so their local gains are defined. In the positive quadrant, the nullclines are $x=f(y)$ and $y=g(x)$, both decreasing. The phase plane arrows point right when $x<f(y)$ and left when $x>f(y)$; they point up when $y<g(x)$ and down when $y>g(x)$. Both axes point into the quadrant. Since $f,g$ are bounded, sufficiently large $x$ or $y$ decreases, giving a bounded forward-invariant rectangle containing all equilibria and eventually containing each trajectory. Thus mutual repression can have one attracting intersection or several intersections separating different basins of attraction.

An equilibrium obeys $x_{\ast }=f(y_{\ast })$, $y_{\ast }=g(x_{\ast })$. Its Jacobian matrix and eigenvalues areThe product is nonnegative. Hence an intersection is a stable equilibrium with two negative real eigenvalues when $f^{\prime }{g}^{\prime }<1$, a saddle equilibrium when $f^{\prime }{g}^{\prime }>1$, and nonhyperbolic when $f^{\prime }{g}^{\prime }=1$. This is the mutual repression stability criterion. To see why an intervening saddle normally accompanies multistability, eliminate $y$ and set $F(x)=f(g(x))-x$. Bounded positivity gives $F(0)>0$ and $F(x)<0$ for large $x$, whileTwo downward, stable crossings require an upward crossing between them. For simple roots that middle crossing has $f^{\prime }{g}^{\prime }>1$ and is a saddle equilibrium. Its stable manifold separates the attracting states. Thus robust multistability requires the loop gain to exceed one at an intervening equilibrium, with the stable equilibria themselves having gain below one. A tangency at gain one is the threshold where equilibria can merge; degenerate crossings require higher-order analysis. A gain exceeding one somewhere is necessary for multiple intersections, but by itself does not prove that a particular pair of nullclines has multiple stable intersections.

Define the logarithmic repression sensitivitiesBoth are negative. At equilibrium $f(y_{\ast })=x_{\ast }$ and $g(x_{\ast })=y_{\ast }$, so their product is exactly $f^{\prime }(y_{\ast })g^{\prime }(x_{\ast })$. The condition can therefore be expressed asEquivalently, the product of the magnitudes of the two negative logarithmic gains must exceed one at that saddle equilibrium. The divergence of the vector field is identically $-2$, so the Bendixson-Dulac criterion excludes periodic orbits in the positive quadrant. The relevant stable long-time states in this bounded phase plane are thus equilibria, rather than attracting limit cycles. The following sketches illustrate the single-attractor and two-attractor possibilities; the saddle in the second panel lies on the diagonal, with its stable diagonal separating the two basins.

For this equal-strength power-law mutual repression, the logarithmic gains and their product at equilibrium areThe mutual repression stability criterion immediately excludes multistability if $mn\le 1$, since $P<mn$ at every finite positive equilibrium. The common prefactor $\lambda$ gives a stronger exclusion: if either exponent is at most one, multistability is impossible for every $\lambda$, including some cases with $mn>1$.

To prove the additional cases, suppose $m\le 1<n$. At equilibrium the two expressions for $\lambda$ implyPut $a=x/(1+x)$ and $b=y/(1+y)$. Because $m\le 1$, the left side is at least $a$, so $a\le b(1-b{)}^{n-1}$. It follows thatThe maximum in the second inequality occurs at $b=2/(n+1)$. Its strict bound follows from $4n/(n+1{)}^2\le 1$, $m\le 1$, and $((n-1)/(n+1){)}^{n-1}<1$. Interchanging the components proves the case $n\le 1<m$; the case with both exponents at most one was already covered by $mn\le 1$. Every root of $F(x)=f(g(x))-x$ therefore has $F^{\prime }<0$. Since $F$ begins positive and ends negative, there is exactly one root: two downward crossings would require another crossing with nonnegative derivative between them. The unique equilibrium is stable.

For completeness, neither exponent can be relaxed above one in this exclusion. If $m>1$ and $n>1$, take $\lambda$ sufficiently large. On $x\in [\lambda /2,\lambda ]$, $g(x)=O({\lambda }^{1-m})\to 0$, so $f(g(x))=\lambda (1+o(1))$ maps that interval into itself. It has a fixed point withIt is a stable equilibrium. By exchanging the components, there is a second stable fixed point with $y\sim \lambda$ and $x\sim {\lambda }^{1-n}$. These two regions are disjoint for large $\lambda$, proving bistability for some production strength. Hence the full answer isFor example, $m=1/2,n=3$ has $mn>1$ but still a unique stable equilibrium for all $\lambda$. Thus the product-only necessary condition for strong feedback is not sufficient when both production prefactors are constrained to be equal.