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# Tripartite Synapse — Biological Reference (companion to v13 pseudocode)
> Companion to `tripartite_synapse_v13_pseudocode.md`. Explains the biology each variable and
> behavior conflates. Variable meanings are unchanged; v13 refined the model logic:
> per-spike calcium deposit (frequency emergent, not a parameter); BEHAVE now separates a FUEL
> shortfall (deposits endurance need) from an OCCUPANCY/structure shortfall (short-term
> depression — no endurance, since more fuel would not help); the endurance feedback term
> (retrograde NO) appears only at the presynapse, where such a signal actually arrives, while
> every other component's endurance proxy is own-state only; EVALUATE merged into TRACE
> (judging a behavior is maintaining a trace); shipment is modelled as a transit-delayed flow
> (axonal/dendritic transport takes a characteristic time, so distal targets receive later),
> reflecting the general principle that every flow has a timescale.
---
## The three synaptic components and their support structures
A SYNAPSE is composed of three first-class components:
- **PRE** — presynaptic bouton (the axon's terminal at this synapse)
- **POST** — postsynaptic spine (the dendrite's terminal at this synapse)
- **ASTRO** — astrosynapse, the perisynaptic astrocytic process (the astrocyte's terminal)
Each has an upstream support structure that supplies it:
- **AXON** supplies PRE (transmission + transport from soma)
- **DEND** supplies POST (integration + transport from soma)
- the **astrocyte cell body** supplies ASTRO (energy + ECM material)
- **SOMA** is the integrating center and the root of neuronal material
The compartment analogy: AXON:PRE = DEND:POST = astrocyte-body:ASTRO = supply line : terminal.
---
## Resource variables
### DAY budget (one per component)
Aggregates fast energy AND fast consumables — everything needed to run moment-to-moment.
- **pre_budget** — ATP for VGCC gating, vesicle fusion (SNARE), VATPase vesicle refill,
plus fast consumables: vesicle membrane lipids, synaptotagmin recycling.
- **post_budget** — ATP for the NaK pump (membrane reset after current), NMDA current
handling, plus fast actin monomers for transient spine changes and receptor-recycling lipids.
- **dend_budget** — ATP for bAP propagation (NaK reset along branch), local translation
(ribosome running cost), SERCA Ca²⁺ resequestration, plus fast mRNA consumed by translation.
- **soma_budget** — ATP for AP generation (Na⁺/K⁺ currents + NaK reset), CREB
phosphorylation, nuclear Ca²⁺ handling, plus shipping running costs.
- **axon_budget** — ATP for AP propagation at nodes of Ranvier, kinesin/dynein motor
running cost, fast myelin maintenance.
- **astro_central_budget** — ATP from glycolysis at the astrocyte cell body; funds EAAT
clearance, serine→D-serine synthesis, lactate export, fast process motility.
### astro_lactate[i]
Lactate exported from the astrocyte cell body to synapse i. Biologically: glucose →
(glycolysis) → lactate, released into extracellular space, absorbed by neuronal MCT2
transporters, converted to pyruvate → TCA → ATP in the neuron's mitochondria. The astrocyte
is the primary fast-energy supplier to pre, post, and dend.
### NIGHT energy (one per component) — NOT recoverable
ATP for structural assembly. Distinct from DAY budget because it is spent on building, and
the work of assembly is thermodynamically gone once done (cannot be recovered by disassembly).
- pre_energy: RIM/Munc13 incorporation, VGCC clustering.
- post_energy: CaMKII anchoring, actin polymerization, PSD scaffold remodeling.
- dend_energy: mitochondria incorporation, cytoskeletal reinforcement.
- soma_energy: ribosome biogenesis, ion-channel incorporation.
- axon_energy: myelination, microtubule stabilization.
- astro_energy: process retraction, ECM secretion, racemase upregulation.
### NIGHT material (one per component) — RECOVERABLE
Slow structural proteins. Recoverable because disassembly (LTD) returns the proteins to a
reusable pool (ubiquitin-proteasome → amino acids; internalized receptors → endosomal reserve).
- **soma_material** (root) — all neuronal structural proteins from CREB-driven synthesis:
AMPA subunits, PSD scaffold, AZ scaffold, mRNA transcripts (Arc, BDNF), organelles.
- **dend_material** — from soma: Arc/plasticity mRNA, mitochondria, cytoskeletal proteins,
AMPA subunits in transit to spines.
- **post_material** — from dend: AMPA receptor subunits (GluA1/2), PSD scaffold (PSD-95,
SHANK, Homer), structural actin, CaMKII.
- **axon_material** — from soma: kinesin/dynein motors, microtubule components, myelin proteins.
- **pre_material** — from axon: RIM, Munc13, VGCC subunits, structural vesicle proteins.
- **astro_material** (root: astrocyte cell body) — EAAT proteins, serine racemase, ECM
proteins (Glypicans, Thrombospondins), process cytoskeleton.
**Why energy and material are separate in NIGHT but combined in DAY:** during DAY both are
fast consumables replenished on the same timescale, so one `budget` variable suffices. During
NIGHT they diverge — material is recoverable after LTD, energy is not — so they must be two
variables. This asymmetry (material returns to the pool, energy is gone) is what makes one
synapse's depression genuinely fund another's potentiation.
---
## Structural variables (strength ceilings — written in NIGHT)
Each aggregates several correlated structural properties into one capacity.
- **pre_structure** — active zone capacity:
slot_ceiling (number of vesicle docking slots) + VGCC_coupling (Ca²⁺-channel proximity to
slots, sets release efficiency) + refill_ceiling (max RRP replenishment rate).
- **post_structure** — spine sensitivity capacity:
slot_ceiling (number of PSD anchoring slots for AMPA) + spine_volume (local reserve and
actin machinery) + reserve_ceiling (endosomal AMPA pool size).
- **dend_structure** — branch capacity:
bAP_fidelity(position) (mitochondrial density sets propagation strength, attenuates with
distance) + translation_ceiling (local mRNA capacity) + transport_speed (cytoskeletal integrity).
- **soma_structure** — somatic output capacity:
baseline_threshold (inverse: ion-channel density at axon initial segment) + AP_reliability
(Na⁺ channel density) + synthesis_ceiling (ribosome density + CREB machinery).
- **axon_structure** — axonal capacity:
propagation reliability (myelination density) + transport_ceiling (motor density + microtubule
integrity) + mitochondrial density.
- **astro_structure** — astrosynaptic environmental capacity:
perisynaptic_distance⁻¹ (wall proximity — closer = more glutamate contained) + EAAT_density
(clearance ceiling) + Dserine_tonic (baseline co-agonist) + ECM_integrity.
**Self-reinforcing both directions:** tighter wrap + more tonic D-serine make future
potentiation easier; looser wrap + zero tonic D-serine make future depression easier.
---
## Budget ceilings (endurance ceilings — written in NIGHT)
- **{component}_budget_ceiling** — the maximum fuel the component can hold / the maximum
duration of sustained behavior. Biologically: mitochondrial density and local fuel-storage
capacity. Built by activity-driven mitochondrial biogenesis; lost by mitophagy when idle.
Parallel to structure: structure is strength capacity, budget_ceiling is endurance capacity.
---
## Trace variables
### fast_trace (one per component) — DAY only, decays automatically
The local record of recent activity that biases the next behavior.
- **pre_fast_trace** — residual presynaptic Ca²⁺ after spikes (τ≈100ms). Biases NT release
(facilitation) and provides tagging eligibility.
- **post_fast_trace** — spine Ca²⁺ amplitude × rise-speed (τ≈tens ms). Encodes the LTP-vs-LTD
instruction (fast rise → CaMKII → potentiation; slow rise → phosphatase → depression).
- **dend_fast_trace** — branch Ca²⁺ from bAP + spine spillover (τ≈300ms). Integrates branch co-activity.
- **soma_fast_trace** — nuclear Ca²⁺ from each AP (τ≈seconds). Drives toward CREB activation.
- **axon_fast_trace** — propagation load (τ≈seconds). High load → Na⁺ inactivation at branch
points → propagation failure (this is axonal short-term depression).
- **astro_fast_trace** — perisynaptic Ca²⁺ from mGluR5 activation by glutamate spillover
(τ≈seconds). Drives D-serine release.
### soma timing traces (emergent refractory + adaptation + alignment)
- **soma_Na_inactivation** (τ≈ms) — sodium-channel inactivation after an AP. Its recovery IS
the refractory period (emergent, not a hardcoded timer). High → absolute refractory; decaying
→ relative refractory; recovered → normal.
- **soma_adaptation** (τ≈100s of ms) — slow K⁺ channel (SK/M-type) activation accumulating
over a spike train, raising threshold. This is spike-frequency adaptation.
- **soma_refractory_alignment** — deposited when a suprathreshold input arrives during
refractoriness (a missed coincidence). Speeds future recovery so the soma aligns to its input
rhythm. Bottom-up: no rhythm is represented; alignment emerges from accumulated local
mismatches and decays when mismatches stop (self-limiting).
### possible_tag (one per component) — intermediate, τ≈smin
Graded accumulation of tagging eligibility. For POST, this is the CANDIDATE tag lifetime.
### endurance_need (one per component) — intermediate, τ≈smin
Deposited when budget depletion interrupts a behavior that was on a LOCALLY successful
trajectory. Records that fuel — not structure, not significance — was the binding constraint
on a forming success. Requires NO dopamine (homeostatic, not associative).
**Local success proxy per component** (each uses only its own state + arrived signals):
- PRE: own fast_trace high (was releasing strongly), optionally amplified by retrograde
messenger (endocannabinoid / NO / BDNF) that has arrived.
- POST: own Ca²⁺ climbing toward tagging threshold (naturally local).
- DEND: own branch strongly active (high branch voltage/Ca²⁺) when propagation fell short.
- SOMA: own nuclear Ca²⁺ climbing toward CREB.
- AXON: own propagation load high (was carrying a strong train).
- ASTRO: own local glutamate/Ca²⁺ high (was under heavy clearance/D-serine demand).
### tag (one per component) — DAY→NIGHT bridge, τ≈hours
The validated record of significance that survives to NIGHT and gates strength commits.
Formed by coincidence of local eligibility + non-local validation (dopamine).
**POST is special — two-phase, three coincidences:**
- CANDIDATE: local Ca²⁺ above threshold + astrosynapse D-serine present (coincidence 1).
- amplified when bAP confirms soma fired (coincidence 2).
- STABLE: CANDIDATE + dopamine within stabilization window (coincidence 3).
Biologically: early CaMKII creates a labile tag (early-LTP); PKA driven by dopamine via D1R
stabilizes it (late-LTP). Without dopamine, the candidate degrades — early-LTP reverses.
---
## Behaviors — biological meaning
### PRE | AP — neurotransmitter release
`NT_flux = RRP × sat(pre_fast_trace, K_release)` models continuous NT release proportional to
the readily-releasable pool and a saturating Ca²⁺ drive (synaptotagmin's cooperative Ca²⁺
sensitivity, simplified to a saturating curve). RRP depletes as released (short-term depression
as a consequence) and refills via VATPase (energy-throttled, so low budget deepens depression).
The mGluR2/3 brake is presynaptic autoinhibition by spillover (Gi → reduced VGCC opening).
### POST | NOT_bAP — three calcium sources, two plasticity cases
- **Source 1 (AMPA):** glutamate opens AMPA → depolarizing current + small Ca²⁺; the
depolarization begins ejecting the NMDA Mg²⁺ block.
- **Source 2 (NMDA):** if depolarized enough (Mg²⁺ ejected) AND D-serine present (astrocyte
co-agonist) AND glutamate bound → large Ca²⁺ influx. This is the coincidence detector.
- **Source 3 (bAP, separate context):** back-propagating AP adds depolarization + Ca²⁺,
amplifying an existing signal supralinearly.
- **Case 1 (STP):** high Ca²⁺ drives AMPA receptors from the local reserve to the surface,
bounded by the anchoring-slot ceiling. Fast, reversible, NO dopamine. When Ca²⁺ falls,
receptors drift back — short-term depression as a passive consequence, never signaled.
- **Case 2 (LTP tag):** high Ca²⁺ + (later) dopamine sets the tag that NIGHT uses to raise the
slot ceiling. NIGHT builds slots; DAY fills them.
### DEND | bAP — bidirectional signaling
Propagates the bAP from soma toward spines (fidelity attenuates with distance — distal spines
get weaker confirmation, are harder to potentiate) and integrates spine signals toward the soma.
### SOMA | AP — integration, firing, emergent timing
Fires when integrated branch input exceeds a threshold that is the baseline (from structure)
raised by adaptation and modulated by neuromodulators, gated by the emergent refractory state.
Each AP deposits three traces (inactivation → refractory, adaptation → threshold rise, nuclear
Ca²⁺ → plasticity). The soma is the coincidence detector at the cellular scale (nuclear Ca²⁺ +
dopamine → CREB), and the production bottleneck: its tag gates how much material all downstream
components get in NIGHT.
### AXON | AP — reliable propagation with frequency-dependent failure
Propagation reliability is set by myelination and degraded by high-frequency load (Na⁺
inactivation at branch points = axonal STD). The axon also transports material to boutons and
sets the timescale of presynaptic structural commits.
### ASTRO | CONTINUOUS — gatekeeper and energy hub
Clears glutamate (EAAT), supplies D-serine (the NMDA co-agonist that gates postsynaptic LTP),
and distributes lactate to the territory by demand-weighting (active synapses generating more
clearance load pull more fuel; slow synapses get less). The same spillover that excites the
astrocyte (mGluR5 → Ca²⁺ → D-serine) also brakes the presynapse (mGluR2/3 → Gi) — one signal,
opposite effects via different receptors. The astrocyte is the energy root and the gain control
of the whole synapse.
---
## NIGHT operations — biological meaning
- **Step 1 (replenish/distribute):** overnight protein synthesis peaks (CREB-driven, gated by
soma_tag — corresponds to slow-wave-sleep replay). Soma material flows to branches/axon then
spines/boutons; astrocyte material flows to astrosynapses, tag-weighted.
- **Step 2 (strength commits):** tagged components build structure — more slots, tighter
coupling, tighter astrosynaptic wrap. Coherence bonus when pre+post+astro all tagged (the
whole synapse agrees). astro_structure self-reinforces.
- **Step 2b (endurance commits):** components with high endurance_need build budget_ceiling —
mitochondrial biogenesis. Competes with step 2 for the same material/energy.
- **Step 3 (passive decay):** both ceilings decay; maintenance from the remaining pool resists
decay only where sufficient. Depotentiation and endurance-loss are both by neglect — no
signal weakens anything; unmaintained capacity simply drifts down. Recovered material (not
energy) returns to pools.
- **Step 4 (homeostatic scaling):** if the soma fired too much overall, all synapses scale down
proportionally (sleep-associated global downscaling), preserving relative differences.
- **Step 5 (clear traces):** fast traces, possible tags, endurance needs, and soma timing traces
reset; tags below expiry clear, above-expiry tags carry forward (multi-night consolidation);
structure and budget_ceiling persist.
### Shockwave lockdown
Emergency global astrocytic Ca²⁺ wave → GABA + ATP release → mass AMPA internalization and
hyperpolarization. Bypasses budget gates. A circuit breaker against runaway excitation.
---
## Pool-filling: private reserve vs contested supply
The pseudocode uses two filling primitives, distinguished by where the resource comes from.
**`fill` (private reserve).** The pool is replenished from a source the component owns
outright, uncontested by siblings, bounded by the component's own ceiling and a rate cap.
- RRP refill — vesicles mobilized from the bouton's own reserve pool toward the docking-slot
ceiling, rate-limited by VATPase. The reserve is private to the bouton.
- SOMA self-replenish — the soma fuels itself from its own mitochondria toward its budget
ceiling. No other component draws on it.
**`refill` (contested supply).** The pool is replenished from a supply that multiple
components compete for, rationed by demand (gap to ceiling).
- pre/post/dend/axon budgets — drawn from astrocytic lactate (shared across all synapses the
astrocyte wraps) plus shipment from soma/axon/dendrite (shared across downstream targets).
**Neither primitive (their own forms).** Some inflows are not fills toward a ceiling:
- AMPA surface insertion — Ca²⁺-driven rate from the spine's private endosomal reserve, with
an explicit passive drift-back (short-term depression) when Ca²⁺ is low. Not a steady fill.
- D-serine release — demand-driven (saturating in astro Ca²⁺) and budget-limited, like NT
release; a release process, not a pool top-up.
- Root productions — `glycolysis(glucose)` at the astrocyte and `CREB_synth(soma_tag)` at the
soma are the system's energy and material roots: raw inflows capped only by the external
vascular supply, not fills toward an internal ceiling.
The distinction matters biologically: a private reserve guarantees a component some autonomy
(the bouton can refill its RRP from its own vesicles even when lactate is scarce), while a
contested supply couples a component's fate to its neighbours' demands (operational budget
fails first where many active synapses compete for the same lactate).
---
## PRE ↔ POST interaction: local computation, message-only coupling
The presynapse and postsynapse never read each other's internal state. They interact only
by writing to and reading from shared cleft channels. Each side computes entirely locally on
what it has: its own variables plus whatever signals have arrived in the cleft. This is the
message-passing realization of the locality principle.
**Forward channel — glutamate (PRE → POST and ASTRO).** The presynapse writes glutamate via
NT_flux. The postsynapse reads it (AMPA, NMDA) and the astrosynapse reads it (clearance,
mGluR5). The astrosynapse clears it. PRE never knows whether POST responded — it only emits.
**Gate channel — astro_Dserine (ASTRO → POST).** The astrosynapse writes D-serine; the
postsynapse reads it as the obligatory NMDA co-agonist. POST cannot open NMDA without this
arrived signal, but it does not read the astrocyte's state — only the delivered D-serine.
**Backward channel + — retro_NO (POST → PRE).** When the postsynapse's NMDA opens (Mg²⁺
ejected, D-serine present, glutamate bound), nNOS — physically tethered to the NMDA receptor
through PSD-95 — synthesises nitric oxide (and, on a slower timescale, BDNF is released).
These diffuse retrogradely to the presynapse. Biologically this is the classic retrograde
messenger of LTP: it tells the bouton that its release landed on a postsynapse that genuinely
responded. In the model, POST emits `retro_NO` proportional to its own NMDA-driven calcium —
computed purely from POST's local state — and PRE reads it as `retro_NO_local`.
`retro_NO_local` is exactly the grounding of the presynaptic endurance signal. The
presynapse's local success proxy is "I was releasing strongly" (`pre_fast_trace` high). On
its own that only says the bouton was working hard, not that the work mattered. `retro_NO`
adds the missing confirmation — that the postsynapse responded — without PRE ever reading
POST's calcium. So PRE deposits endurance need as `pre_fast_trace × (1 + retro_NO_local)`:
strong release that was confirmed effective makes the strongest claim that fuel, not
futility, was what interrupted a forming success. retro_NO is short-lived (NO degrades and
diffuses within seconds), so the channel decays fast — confirmation must be recent to count.
**Backward channel — retro_eCB (POST → PRE).** When the postsynapse is strongly
depolarised, it synthesises endocannabinoids (2-AG, anandamide) that diffuse retrogradely and
bind presynaptic CB1 receptors, suppressing release. This is depolarisation-induced
suppression of excitation (DSE) — a homeostatic negative feedback: an over-driven postsynapse
tells the presynapse to release less. In the model, POST emits `retro_eCB` from its own
membrane potential, and PRE reads it as `retro_eCB_local`, which reduces the release drive
`sat(...) × (1 - retro_eCB_local)`. Again POST computes from its own state; PRE adjusts from
the arrived signal; neither reads the other's interior.
The two backward channels are opposite-signed messages the postsynapse sends about its own
condition: retro_NO says "your input was effective — worth sustaining," retro_eCB says "I am
saturated — ease off." Together with the forward glutamate and the D-serine gate, they make
the synapse a fully message-coupled system of locally-computing components.
**Why RRP refill is in NOT_AP only.** During an AP the bouton releases — RRP depletes. Refill
(VATPase reloading vesicles from the reserve pool) is a recovery process that proceeds between
spikes. Placing `fill(RRP, ...)` only in the NOT_AP context makes the AP context pure
depletion and the NOT_AP context pure recovery. A consequence falls out for free: during
sustained high-frequency firing there are many AP steps and few NOT_AP steps, so RRP depletes
faster than it recovers — short-term depression deepens with frequency, with no explicit
depression rule. The release itself is throttled further when budget is low (VATPase refill
is energy-limited), coupling metabolic state to the depth of depression.
@@ -0,0 +1,502 @@
# Tripartite Synapse — Pseudocode v13
> Companion: `tripartite_synapse_v13_biology.md`.
> Changes from v12:
> (1) per-spike calcium deposit (frequency stays emergent, not a parameter)
> (2) BEHAVE splits FUEL shortfall (→ endurance) from OCCUPANCY shortfall (→ STD)
> (3) endurance feedback term only where a retrograde signal actually arrives
> (4) TRACE absorbs EVALUATE → seven-group grammar
> (5) parameter declarations completed (ship_cost, ACh_gain, NIGHT block)
> (6) shipment is a transit-delayed flow; every flow has a timescale
> (7) all group labels are standalone headers at one column
---
## Functional groups (seven-group grammar)
```
RECEIVE take in resources + signals that arrived from outside (boundary: in)
TRACE maintain the trace hierarchy — deposit fast trace; accumulate
possible_tag + endurance_need; stabilize tag on coincidence
ADJUST compute local operating parameters from structure + traces + modulators
BEHAVE the component's defining action, within both ceilings
EMIT send out — signals (messages) + resources (shipments) (boundary: out)
RECOVER refill own private pools consumed by behaving
DECAY let traces recede, closing their windows
```
EVALUATE merged into TRACE: judging a behavior is always maintaining a trace, whether or not
a trace is written. BEHAVE and EMIT stay separate — EMIT is the output half of the locality
interface (RECEIVE/EMIT are the only boundary crossings). TRACE spans all timescales: the
soma's inactivation, adaptation, and nuclear-Ca deposits are all TRACE. Order within a context
follows data dependencies; TRACE reads/writes whatever trace state is current.
EVERY FLOW HAS A TIMESCALE. Decay relaxes toward 0 over τ; creation/arrival relaxes toward a
target over τ — the same first-order operator. Within-step writes are the special case τ ≪ Δt.
Rate-limited inflows (fill/refill/flux·Δt) carry their τ implicitly; shipment carries an
explicit transit delay (see `transit`).
---
## Conventions
```
SCOPE = {DAY, NIGHT} CONTEXT = {AP, NOT_AP, bAP, NOT_bAP, CONTINUOUS}
DAY budget · fast_trace · possible_tag · endurance_need
BRIDGE tag (POST: CANDIDATE→STABLE)
NIGHT energy (not recoverable) · material (recoverable) · structure · budget_ceiling
LOCALITY only local state + arrived signals; no component reads another's internal state.
CLEFT MESSAGE CHANNELS SHIPMENT CHANNELS (transit-delayed)
glutamate PRE → POST, ASTRO soma_ship_dend SOMA→DEND
astro_Dserine ASTRO → POST soma_ship_axon SOMA→AXON
retro_NO POST → PRE (+) dend_ship_post DEND→POST
retro_eCB POST → PRE () axon_ship_pre AXON→PRE
```
---
## Primitives (return the increment; caller applies it)
```
sat(x, K) = x / (K + x)
fill(pool, ceiling, rate, cost, budget) -> amount: // PRIVATE reserve, rate-limited (implicit τ)
amount = min(rate, ceiling - pool)·Δt; budget -= amount·cost; return amount
refill(c from supply S) -> amount: // CONTESTED supply, gap-bounded
demand = c.budget_ceiling - c.budget
factor = min(1, S / (Σ demand over components on S + ε)); S -= demand·factor
return demand·factor
ship(from_budget, demand_sig, frac, cost) -> amount: // emit into transit (not to target directly)
amount = min(from_budget·frac, demand_sig); from_budget -= amount·(1+ship_cost); return amount
transit(channel, τ_transport) -> arrival: // delivers in-transit cargo over τ
arrival = channel·(Δt/τ_transport); channel -= arrival; return arrival
```
---
## SHARED parameters
```
dopamine NE ACh // organism broadcasts (external)
glucose geometry // physical (external)
elig dop_thr tag_thr tag_expiry // strength gates (universal)
traj_thr endur_thr // endurance gates (universal)
ship_cost // transport overhead (all shipments)
τ_transport_{dend,axon,spine,bouton} // shipment transit times (distance-dependent)
ε
```
## NIGHT parameters (consolidation only)
```
slot_cost cap_cost f_cap // commit sizes / endurance fraction
maint_frac cap_frac // maintenance allocation
decay_rate capacity_decay_rate recycle // passive decay + recovery
homeostatic_ceiling coherence_factor assembly_cost biogenesis_cost maint_cost
f_dend f_axon f_spine f_bouton // material distribution fractions
{dend,axon,pre,post}_ship_frac // DAY shipment fractions
{dend,axon,pre,post}_energy_frac // energy distribution fractions
```
---
---
# DAY
---
## PRE
```
// PARAMETERS K_release · release_cost · fusion_cost · vatpase_cost · spillover · brake
// INTERFACE
// EMIT glutamate → POST, ASTRO
// RECEIVE astro_lactate[syn] ← ASTRO ; axon_ship_pre ← AXON ; retro_NO, retro_eCB ← POST
// pre_material ← AXON(NIGHT) ; pre_energy ← SOMA(NIGHT)
// READ glutamate (own cleft, autobrake) ; dopamine (gates tag)
// OWN pre_structure{slot_ceiling, VGCC_coupling, refill_ceiling} ; pre_budget_ceiling
// EMERGENCY shockwave_lockdown ← ASTRO
DAY | AP:
// TRACE (Ca²⁺ bolus from THIS spike — also drives release; frequency is emergent)
pre_fast_trace += spike_Ca(pre_structure.VGCC_coupling)
// ADJUST (release drive from residual Ca²⁺ + received DSE brake)
drive = sat(pre_fast_trace, K_release) × (1 - retro_eCB_local)
// BEHAVE (release; two distinct failure modes)
if pre_budget < release_cost:
// FUEL shortfall → endurance evidence (retro_NO-confirmed local success)
suppress(NT_flux)
if pre_fast_trace > traj_thr:
pre_endurance_need += pre_fast_trace × (1 + retro_NO_local)
exit
if RRP == 0:
// OCCUPANCY shortfall → short-term depression (NOT endurance; fuel was fine)
suppress(NT_flux)
exit
NT_flux = RRP × drive; RRP -= NT_flux·Δt; pre_budget -= NT_flux·fusion_cost
// EMIT (glutamate into cleft)
glutamate += NT_flux·Δt
if glutamate > spillover: drive *= brake // own-cleft autobrake
DAY | NOT_AP:
// RECEIVE (latch backward messages ; replenish budget from contested supply)
retro_NO_local = retro_NO; retro_eCB_local = retro_eCB
pre_budget += refill(pre from astro_lactate[syn] + transit(axon_ship_pre, τ_transport_bouton))
// TRACE (strength: eligibility → tag via dopamine)
if pre_fast_trace > elig: pre_possible_tag += pre_fast_trace
if dopamine > dop_thr and pre_possible_tag > tag_thr:
pre_tag += dopamine × pre_possible_tag
// RECOVER (RRP from private reserve toward its ceiling)
RRP += fill(RRP, pre_structure.slot_ceiling, pre_structure.refill_ceiling, vatpase_cost, pre_budget)
// DECAY
pre_fast_trace *= decay(100ms); pre_possible_tag *= decay(s)
pre_endurance_need *= decay(min); pre_tag *= decay(hr)
dopamine *= decay(ms); retro_NO *= decay(s); retro_eCB *= decay(s)
```
---
## POST
```
// PARAMETERS K_AMPA · AMPA_Ca · AMPA_cost · NMDA_cost · bAP_cost · pka_cost · traffic_cost
// req_cost · Mg_eject · Dserine_thr · Ca_STP · Ca_TAG · eCB_thr · drift · baseline
// NO_synth_cost · eCB_synth_cost
// INTERFACE
// EMIT retro_NO (+), retro_eCB () → PRE
// RECEIVE astro_lactate[syn] ← ASTRO ; dend_ship_post ← DEND ; post_material ← DEND(NIGHT) ; post_energy ← SOMA(NIGHT)
// READ glutamate ← PRE ; astro_Dserine ← ASTRO ; bAP (dend_structure.bAP_fidelity) ; dopamine
// OWN post_structure{slot_ceiling, spine_volume, reserve_ceiling} ; post_budget_ceiling
// EMERGENCY shockwave_lockdown ← ASTRO
// NOTE POST endurance is own-state only (own Ca climbing); no arrived feedback term.
DAY | NOT_bAP:
// RECEIVE
post_budget += refill(post from astro_lactate[syn] + transit(dend_ship_post, τ_transport_spine))
// ADJUST (AMPA drive from arrived glutamate)
a = sat(glutamate, K_AMPA)
// BEHAVE (SOURCE 1 AMPA: current + small Ca + begins Mg ejection)
AMPA_current = a × AMPA_surface; Vm += AMPA_current; post_budget -= AMPA_cost
// TRACE (Ca deposited by AMPA)
post_fast_trace += AMPA_Ca·AMPA_current
// BEHAVE (SOURCE 2 NMDA: large Ca on local coincidence)
if Vm > Mg_eject and astro_Dserine > Dserine_thr and glutamate > 0:
post_fast_trace += NMDA_Ca(glutamate)·rise_speed(); post_budget -= NMDA_cost
// EMIT (+ NO/BDNF: "release reached a responsive target")
retro_NO += NO_emit(post_fast_trace); post_budget -= NO_synth_cost
// EMIT ( endocannabinoid / DSE when over-driven)
if Vm > eCB_thr:
retro_eCB += eCB_emit(Vm); post_budget -= eCB_synth_cost
post_fast_trace *= decay(ms)
// BEHAVE (STP fill slots from private reserve ; else STD drift = consequence)
if post_fast_trace > Ca_STP:
if post_budget < traffic_cost:
// FUEL shortfall → endurance (own Ca was climbing toward a tag)
if post_fast_trace > traj_thr and post_fast_trace_rising:
post_endurance_need += post_fast_trace
else if AMPA_surface < post_structure.slot_ceiling:
AMPA_surface += Ca_insert(post_fast_trace); post_budget -= traffic_cost
// else: surface already at slot_ceiling → structure-limited (not endurance)
else:
AMPA_surface = max(AMPA_surface - drift·Δt, baseline) // STD = consequence
// TRACE (strength: CANDIDATE then STABLE via dopamine)
if post_fast_trace > Ca_TAG: post_possible_tag += post_fast_trace; post_budget -= pka_cost
if dopamine > dop_thr and post_possible_tag > tag_thr:
post_tag += dopamine × post_possible_tag
// DECAY
post_possible_tag *= decay(min); post_endurance_need *= decay(min)
post_tag *= decay(hr); dopamine *= decay(ms)
DAY | bAP:
// BEHAVE (SOURCE 3 bAP: depolarization + Ca, amplifies existing signal)
Vm += bAP_depol × dend_structure.bAP_fidelity; post_budget -= bAP_cost
// TRACE (supralinear boost only if a CANDIDATE is present)
if post_possible_tag > Ca_TAG: post_fast_trace += bAP_Ca_boost()
```
---
## DEND
```
// PARAMETERS prop_cost · branch_Ca_cost · integrate_cost · translate_cost · ACh_gain
// INTERFACE
// EMIT bAP_local → POST ; branch_Vm → SOMA ; dend_ship_post → POST
// RECEIVE astro_lactate[branch] ← ASTRO ; soma_ship_dend ← SOMA ; dend_material, dend_energy ← SOMA(NIGHT)
// READ SOMA.fired ; POST.Vm + spine spillover ; dopamine ; ACh
// OWN dend_structure{bAP_fidelity(pos), translation_ceiling, transport_speed} ; dend_budget_ceiling
// NOTE DEND endurance fires only on FUEL-limited propagation, not structural attenuation;
// own-state proxy (strong branch activity); no arrived feedback term.
DAY | bAP:
// ADJUST (propagation strength from structure — inside propagate())
// BEHAVE (propagate bAP; distinguish fuel-limited vs structure-limited shortfall)
if dend_budget < prop_cost:
// FUEL shortfall → endurance (branch was strongly active)
if dend_fast_trace > traj_thr:
dend_endurance_need += dend_fast_trace
bAP_local, reached = propagate_partial(dend_budget)
else:
bAP_local, reached = propagate(SOMA.fired, dend_structure.bAP_fidelity, geometry)
// reached < full here is structural attenuation (distance), NOT endurance
dend_budget -= prop_cost × reached
// TRACE
dend_fast_trace += bAP_Ca(bAP_local) + spine_spillover(); dend_budget -= branch_Ca_cost
// EMIT (integrated voltage to soma ; propagated bAP already reached spines)
branch_Vm = integrate(POST.Vm, spines); dend_budget -= integrate_cost
DAY | NOT_bAP:
// RECEIVE
dend_budget += refill(dend from astro_lactate[branch] + transit(soma_ship_dend, τ_transport_dend))
// TRACE (strength)
if dend_fast_trace > elig: dend_possible_tag += dend_fast_trace
if dopamine > dop_thr and dend_possible_tag > tag_thr:
dend_tag += dopamine × dend_possible_tag
// ADJUST (commit threshold lowered by attention)
commit_threshold *= 1/(1 + ACh·ACh_gain)
// EMIT (ship budget to spines; demand = post tag)
dend_ship_post = ship(dend_budget, post_demand, post_ship_frac, ship_cost)
// BEHAVE (local translation if tagged — fills dend capacity faster)
if dend_tag > tag_expiry and dend_budget > translate_cost: dend_budget -= translate_cost
// DECAY
dend_fast_trace *= decay(300ms); dend_possible_tag *= decay(s)
dend_endurance_need *= decay(min); dend_tag *= decay(hr)
```
---
## SOMA
```
// PARAMETERS ap_cost · nuclear_cost · creb_cost · mito_output · inactivation · ap_amp · ap_contrib
// base_recovery · τ_Na · τ_adapt · τ_nuclear · τ_align
// INTERFACE
// EMIT fired → AXON (propagate) + DEND (bAP) ; soma_ship_dend → DEND ; soma_ship_axon → AXON
// RECEIVE self (mitochondria, ROOT) ; branch_Vm ← DEND
// READ dopamine ; NE ; ACh
// OWN soma_structure{baseline_threshold, AP_reliability, synthesis_ceiling} ; soma_budget_ceiling
// NOTE SOMA endurance fires only on FUEL shortfall (budget < ap_cost);
// refractory / sub-threshold are timing limits, not endurance. Own-state proxy.
DAY | AP:
// ADJUST (threshold from structure + adaptation + neuromodulators ; refractory gate)
threshold = soma_structure.baseline_threshold × (1 + soma_adaptation) × neuromod(NE, ACh)
can_fire = soma_Na_inactivation < inactivation
// BEHAVE (fire if able)
if branch_Vm > threshold and can_fire:
if soma_budget < ap_cost:
// FUEL shortfall → endurance (firing was approaching CREB)
if soma_fast_trace > traj_thr and soma_fast_trace_rising:
soma_endurance_need += soma_fast_trace
exit
// EMIT (fired → AXON, DEND)
fired = True; soma_budget -= ap_cost
// TRACE (three traces from one AP — all timescales)
soma_Na_inactivation += ap_amp // → refractory (emergent)
soma_adaptation += ap_contrib // → threshold rise
soma_fast_trace += nuclear_Ca(); soma_budget -= nuclear_cost
// TRACE (strength)
if soma_fast_trace > elig: soma_possible_tag += soma_fast_trace
if dopamine > dop_thr and soma_possible_tag > tag_thr:
soma_tag += dopamine × soma_possible_tag
soma_budget -= creb_cost
DAY | NOT_AP:
// RECEIVE (self-replenish from private root ; integrate input)
soma_budget += fill(soma_budget, soma_budget_ceiling, mito_output, 0, soma_budget)
branch_Vm = integrate(DEND.branch_Vm, branches)
// TRACE (bottom-up refractory alignment: suprathreshold input during refractory)
if branch_Vm > threshold and soma_Na_inactivation > inactivation:
soma_refractory_alignment += (branch_Vm - threshold) × soma_Na_inactivation
// EMIT (ship downstream into transit; demand = propagated tags)
soma_ship_dend = ship(soma_budget, dend_demand, dend_ship_frac, ship_cost)
soma_ship_axon = ship(soma_budget, axon_demand, axon_ship_frac, ship_cost)
// RECOVER (inactivation recovery sped by alignment trace → emergent refractory)
recovery = base_recovery × (1 + soma_refractory_alignment)
soma_Na_inactivation *= decay(τ_Na / recovery)
// DECAY
soma_adaptation *= decay(τ_adapt); soma_fast_trace *= decay(τ_nuclear)
soma_refractory_alignment *= decay(τ_align) // self-limiting
soma_possible_tag *= decay(s); soma_endurance_need *= decay(min)
soma_tag *= decay(hr); dopamine *= decay(ms)
```
---
## AXON
```
// PARAMETERS prop_cost · budget_factor
// INTERFACE
// EMIT APs_delivered → PRE (propagation) ; axon_ship_pre → PRE
// RECEIVE soma_ship_axon ← SOMA ; astro_lactate[shaft] ← ASTRO ; axon_material, axon_energy ← SOMA(NIGHT)
// READ SOMA.fired ; dopamine
// OWN axon_structure{propagation, transport_ceiling, mito_density} ; axon_budget_ceiling
// NOTE AXON endurance fires only on FUEL shortfall; load-driven failure fail(fast_trace)
// is axonal STD (a consequence), not endurance. Own-state proxy.
DAY | AP:
// ADJUST (reliability from structure load-driven failure)
reliability = axon_structure.propagation × (1 - fail(axon_fast_trace)) // fail() = STD, not endurance
// BEHAVE (propagate; FUEL shortfall degrades + flags endurance)
if axon_budget < prop_cost:
reliability *= budget_factor
if axon_fast_trace > traj_thr: // FUEL-limited → endurance
axon_endurance_need += axon_fast_trace
delivered = fired × reliability; axon_budget -= prop_cost × delivered
// EMIT (delivered APs reach boutons)
// TRACE
axon_fast_trace += delivered; axon_fast_trace *= decay(s)
DAY | NOT_AP:
// RECEIVE
axon_budget += refill(axon from soma_ship_axon + astro_lactate[shaft])
// TRACE (strength)
if axon_fast_trace > elig: axon_possible_tag += axon_fast_trace
if dopamine > dop_thr and axon_possible_tag > tag_thr:
axon_tag += dopamine × axon_possible_tag
// EMIT (ship to boutons; demand = pre tag)
axon_ship_pre = ship(axon_budget, pre_demand, pre_ship_frac, ship_cost)
// DECAY
axon_fast_trace *= decay(s); axon_possible_tag *= decay(s)
axon_endurance_need *= decay(min); axon_tag *= decay(hr)
```
---
## ASTRO
```
// PARAMETERS K_Dserine · Ds_max · Ds_frac · Ds_cost · EAAT_cost · lactate_cost · spillover · overload
// INTERFACE
// EMIT astro_lactate[i] → pre/post/dend budgets ; astro_Dserine[i] → POST (gate)
// RECEIVE glucose (ROOT) ; astro_material, astro_energy ← cell body (NIGHT)
// READ glutamate ← PRE (clearance + spillover) ; dopamine
// OWN astro_structure{perisynaptic_distance⁻¹, EAAT, Dserine_tonic, ECM} ; astro_budget_ceiling
// NOTE ASTRO endurance fires on BUDGET-limited synthesis (got<want via low budget);
// material/precursor-limited synthesis is a material limit, not endurance. Own-state proxy.
// EMERGENCY emits shockwave_lockdown on overload
DAY | CONTINUOUS: // per astrosynapse i
// RECEIVE (root production, capped by glucose)
astro_central_budget += glycolysis(glucose)·Δt
// ADJUST (demand weights across territory)
for each i: demand[i] = clearance_load[i] × astro_structure[i].delivery_eff
factor = min(1, astro_central_budget / (Σ demand·lactate_cost + ε))
// EMIT (demand-weighted lactate to all components)
for each i:
astro_lactate[i] = demand[i] × factor; astro_central_budget -= astro_lactate[i]·lactate_cost
// BEHAVE (clear glutamate ; supply tonic D-serine)
glutamate[i] -= astro_structure[i].EAAT × glutamate[i]·Δt; astro_central_budget -= clearance·EAAT_cost
astro_Dserine[i] += astro_structure[i].Dserine_tonic·Δt
if glutamate[i] > spillover:
// TRACE
astro_fast_trace[i] += mGluR_Ca(); astro_fast_trace[i] *= decay(s)
// ADJUST (D-serine demand from spillover)
want = sat(astro_fast_trace[i], K_Dserine) × Ds_max
got = min(want, astro_central_budget × Ds_frac)
// BEHAVE + EMIT (D-serine pulse to POST gate)
astro_Dserine[i] += got; astro_central_budget -= got·Ds_cost
// TRACE (endurance: BUDGET-limited synthesis under high own demand)
if got < want and astro_central_budget low and astro_fast_trace[i] > traj_thr:
astro_endurance_need[i] += (want - got)
// TRACE (strength)
if astro_fast_trace[i] > elig: astro_possible_tag[i] += astro_fast_trace[i]
if dopamine > dop_thr and astro_possible_tag[i] > tag_thr:
astro_tag[i] += dopamine × astro_possible_tag[i]
// DECAY
astro_possible_tag[i] *= decay(s); astro_endurance_need[i] *= decay(min); astro_tag[i] *= decay(hr)
// EMERGENCY
if astro_fast_trace[i] > overload: emit(shockwave_lockdown)
```
---
## Special — Shockwave Lockdown
```
DAY or NIGHT | OVERLOAD:
Vm = HYPERPOLARIZED; AMPA_surface = mass_internalize() → post reserve
axon_fast_trace += overdrive(); astro_central_budget -= emergency_cost
```
---
---
# NIGHT
Same grammar on ceilings, once per cycle.
```
NIGHT | 1 RECEIVE + ADJUST + EMIT (replenish, weight, distribute)
// RECEIVE overnight production at roots (capped by glucose; gated by soma_tag)
astro_central_{budget,energy,material} += overnight_*(glucose, …)·Δt
soma_{budget,energy} += overnight_*()·Δt ; soma_material += CREB_synth(soma_tag)·Δt
// ADJUST tag-weighted shares
for each i with astro_tag[i] > tag_expiry: w = astro_tag[i]/Σastro_tag
// EMIT distribute material + energy down the supply chains
astro_energy[i] += astro_central_energy·w; astro_material[i] += astro_central_material·w
dend_material += soma_material·f_dend ; axon_material += soma_material·f_axon ; soma_material -= …
post_material += dend_material·f_spine ; pre_material += axon_material·f_bouton
{pre,post,dend,axon}_energy += soma_energy·{·}_energy_frac
{pre,post,dend,axon}_budget += astro_lactate_reserve·{·}_frac·Δt
NIGHT | 2 TRACE + BEHAVE (strength commits → raise structure)
// ADJUST coherence bonus when pre, post, astro tags align
coherence = (pre_tag, post_tag, astro_tag all > tag_expiry) ? coherence_factor : 1
// TRACE+BEHAVE build structure for tagged components
for each c with c_tag > tag_expiry:
Δ = min(slot_cost, c_material, c_energy·f_cap)
c_structure += Δ × (coherence if c in {pre,post,astro} else 1)
c_material -= Δ; c_energy -= Δ·assembly_cost; if Δ < slot_cost: queue(→ next NIGHT)
NIGHT | 2b TRACE + BEHAVE (endurance commits → raise budget_ceiling ; no dopamine ; competes w/ 2)
for each c with c_endurance_need > endur_thr:
Δ = min(cap_cost, c_material·f_cap, c_energy·f_cap)
c_budget_ceiling += Δ; c_material -= Δ; c_energy -= Δ·biogenesis_cost; if Δ<cap_cost: queue
NIGHT | 3 DECAY + RECOVER (both ceilings decay by neglect ; material recovered)
maint = (total_material - consumed) × maint_frac / synapse_count
for each synapse: // DECAY
{pre,post,dend,astro}_structure -= decay_rate·Δt
{pre,post,dend,astro}_budget_ceiling -= capacity_decay_rate·Δt
if maint ≥ maint_cost: structure += full_maint ; budget_ceiling += full_cap_maint
else: structure += maint·frac ; budget_ceiling += maint·cap_frac
for each synapse with net_change < 0: // RECOVER
{pre,post,astro}_material += |net_change|·recycle·frac // material recovered, energy not
NIGHT | 4 BEHAVE (homeostatic scaling)
if soma_tag > homeostatic_ceiling:
s = homeostatic_ceiling / soma_tag
for each synapse: post_structure.slot_ceiling *= s ; pre_structure.slot_ceiling *= s
soma_material += Σ reduction·recycle
NIGHT | 5 DECAY (clear traces)
all fast_trace, possible_tag, endurance_need = 0
soma_Na_inactivation = soma_adaptation = soma_refractory_alignment = 0
for each tag: if tag < tag_expiry: tag = 0 // else carry forward
// structure and budget_ceiling PERSIST
```
---
## One-view summary
```
SEVEN-GROUP GRAMMAR, TWO TIMESCALES
RECEIVE · TRACE · ADJUST · BEHAVE · EMIT · RECOVER · DECAY
DAY grammar on OCCUPANCY within two ceilings (structure=strength, budget_ceiling=endurance)
TRACE yields two evidence streams from local state + arrived signals:
fast_trace + dopamine → tag (strength)
FUEL shortfall + interrupted LOCAL success → endurance_need (endurance)
OCCUPANCY/structure/timing shortfalls → short-term depression (NOT endurance)
NIGHT same grammar on the CEILINGS:
tag → structure ; endurance_need → budget_ceiling ; both draw one pool (compete)
unmaintained ceilings decay → recovered material funds the rest
FLOWS every flow has a timescale: decay relaxes toward 0 over τ, arrival toward a target
over τ; shipment is transit-delayed; rate-limited inflows carry τ implicitly.
LOCAL every group uses only own state + arrived signals; RECEIVE/EMIT are the only crossings.
```