
the study proceeds.
The second element of the model captures the integra-
tion and leak of the current generated when the SQUID is
driven above threshold. These behaviors are accomplished
by adding an L-Rloop to the output of the SQUID, as
shown by the integration loop labeled I in Fig. 2(b). The
current integrated in this branch of the circuit is the quan-
tity of interest for the dendrite. It is this quantitiy that
will couple to other dendrites or the neuron cell body, and
it is this quantity we wish to track with our phenomenolog-
ical model. Dendrites comprising a receiving loop coupled
to an integrating loop [Fig. 2(b)] are referred to as RI den-
drites. We know from elementary circuit theory that the
L-Rloop will result in exponential decay of signal with a
time constant of the dendritic integration loop given by
τdi =Ldi/Rdi.
We can now write down a postulated expression for the
signal sstored in the integration loop of a dendrite:
βd s
dτ =r(φ, s;ib)−α s. (2)
Equation 2 states that the signal sgrows in time due to the
driving term, which is the rate of flux quantum produc-
tion, denoted by r. This function rdepends on the applied
flux to the receiving loop of the dendrite, φ= Φ/Φ0, as
well as of the signal present in the integration loop, s. The
rate also depends on the bias current, ib, as a parameter
that throughout this work is assumed to be held constant
over times much longer than the inter-fluxon interval. In
Eq. 2 we have formulated the model in dimensionless units,
where s=Idi/Ic, and Idi is the current present in the
dendritic integration loop. β= 2 π IcLdi/Φ0is a dimen-
sionless parameter that quantifies the inductance of the
loop, and α=Rdi/rj, where rjis the shunt resistance of
each JJ in the resistively and capacitively shunted junc-
tion model [20–23]. The signal sdecays at a rate related
to αand β. Specifically, the time constant for decay is
given by τdi =Ldi/Rdi =β/ωcα, where ωcis the Joseph-
son characteristic frequency discussed in Appendix A, and
we include the subscript on τdi to refer to the dendritic
integration loop and distinguish that quantity from the
dimensionless time variable entering Eq. 2. Equation 2 is
a leaky integrator ODE. The drive term is the rate of flux
quantum production, and the leak term gives simple ex-
ponential decay, as expected from an L-Rcircuit.
Dendrites are coupled to each other through flux. The
coupling flux from dendrites indexed by jto dendrite iis
given by
φi=
n
X
j=1
Jij sj,(3)
where Jij is a coupling term proportional to the mutual
inductance that includes contributions from all the trans-
formers present on the collection coil in Fig. 2(b). Equa-
tion 3 shows that coupling between dendrites is due to the
signal in the integration loop of one dendrite being com-
municated as flux into the receiving loop of a subsequent
dendrite. The signal in the subsequent dendrite is then
obtained through the evolution of Eq. 2 with the flux from
the first dendrite providing the flux φin the function r
and the signal from the second dendrite providing the s
term. The ibterm entering rrefers to the bias on the
second dendrite and is treated here as a parameter rather
than a dynamical variable. Further details regarding the
derivation of these expressions is given in Appendix C.
Equations 2 and 3 constitute the phenomenological
model of a dendrite. A neuron or network can be sim-
ulated by solving these coupled equations for all dendrites
in the system. However, we have not specified the form
for the rate function, r(φ, s;ib), which is central to the
model.
We have arrived at Eq. 2 as a postulate; this expres-
sion is not directly obtained from the underlying circuit
equations. The postulate is that there will be a function
r(φ, s;ib) such that Eq. 2 provides an accurate account of
the signal present in a dendrite’s integration loop under
the circumstances of interest for loop neurons, provided
we only inquire about the signal over time scales appre-
ciably longer than the inter-fluxon interval, which is on
the order of 10 ps. We aim to interrogate dendrites on
time scales of 100 ps or longer, with neuron and network
activity of interest on time scales from nanoseconds to the
longest timescales that can be simulated under the limits
of computational resources.
Our procedure for obtaining r(φ, s;ib) is as follows. A
dendrite with a SQUID as a receiving loop and an L-
Rbranch as an integrating loop is numerically modeled
with the circuit equations given in Appendix A. A con-
stant value of flux is applied to the receiving loop, and the
rate of flux-quantum production is monitored as a func-
tion of time while current accumulates in the integration
loop. For these simulations, the resistance of the integra-
tion loop is set to zero. This procedure is repeated for
many values of φand ibto obtain what we refer to as
“rate arrays”, which are shown in Fig. 4, where r(φ, s;ib)
is plotted as a function of φand sfor three values of ib.
Here we work in dimensionless units, so the units of r
are fluxons generated per unit of dimensionless time, τ,
which is related to the JJ characteristic frequency (Ap-
pendix A). It can be seen that the value of r(φ, s;ib) is
monotonically increasing with φover the range considered
here, while accumulation of sdecreases the rate of flux-
quantum production. This decrease is because addition of
current to the integration loop diverts the bias away from
the SQUID, so the voltage is decreased, and the rate is
reduced in accordance with Eq. 1. When sufficient signal
is accumulated in the integration loop, the rate of flux
quantum production drops to zero, and we say the loop is
saturated.
For a small value of ib[Fig. 4(a)] a large amount of flux is
required to drive the dendrite above threshold to the active
state, and a small signal sis present at saturation. As ibis
increased [Figs. 4(b) and (c)] the threshold is reduced, and
the saturation level is increased. The qualitative shape of
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