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ocrampal
2026-04-01 12:03:56 +02:00
parent 537ac9428c
commit 865f3952da
4 changed files with 321 additions and 95 deletions
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neuron/BEH-AXO.md
+5 -5
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@@ -198,7 +198,7 @@ container: BEH-PRE
```
### ms: behaviors
### ms: behaviors PRE
#### AP-RRPConcentration: Context
@@ -402,14 +402,14 @@ episode: CaTracesAccumulationSlow
trace: None
```
### sec: behaviors
### sec: behaviors PRE
#### SecContext: Context
#### CheckConditions: Context
Contestualizziamo in maniera Fixed ogni mezzo secondo?
```Gen
context: SecContext
context: CheckConditions
contained_by: BEH-PRE
in_context: Fixed
@@ -540,7 +540,7 @@ episode: RPShuttleMaximal
trace: None
```
### min: behaviors
### min: behaviors PRE
#### Refill RP from Glutamine
neuron/BEH-BD.md
+38 -22
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@@ -15,7 +15,7 @@ Qui BEH-DB espande solo i BEH-POST, e' un cavo di collegamento come l'assone
Container: BEH-BD
expansion:
- BEH-POST ( fullness: 50x, active: 20x, emptiness: 10x )
- BEH-POST ( full: 50x, active: 20x, empty: 10x )
modulated_by: DEV-BD-BEH-POST-TUB from DEV-N.md
```
@@ -27,7 +27,7 @@ Like its presynaptic partner, the postsynapse is governed by three interlocking
---
### 1. The $V_{post}$ Loop: The Fast Gatekeeper (Milliseconds)
1.The $V_{post}$ Loop: The Fast Gatekeeper (Milliseconds)
This is the primary electrophysiological response, where chemical signals are converted back into electricity.
@@ -39,7 +39,7 @@ This is the primary electrophysiological response, where chemical signals are co
---
### 2. The $Ca^{2+}$ Loop: The Plasticity Controller (Seconds)
2.The $Ca^{2+}$ Loop: The Plasticity Controller (Seconds)
This loop translates electrical timing into biological "memory."
@@ -51,7 +51,7 @@ This loop translates electrical timing into biological "memory."
---
### 3. The ATP Loop: The Metabolic Backbone (Minutes)
3.The ATP Loop: The Metabolic Backbone (Minutes)
This is the "Hidden Master" that determines if the other two loops are allowed to function.
@@ -71,36 +71,46 @@ The system is beautifully asymmetric. While the presynapse is built to **supply*
container: BEH-POST
expansion:
- BEH-POST-AMPA ( fullness: 10x, active: 5x, emptiness: 2x )
- BEH-POST-AMPA ( full: 10x, active: 5x, empty: 2x )
# modulated_by: TUN-POST-IC # possible/actual
tub_local:
- Ca2+ ( fullness: 60x, active: 30x, emptiness: 0x )
- Ca2+ ( full: 60x, active: 30x, empty: 0x )
# modulated_by: DEV-POST-???-FULL # Full
- Nox ( fullness: 100x, active: 20x, emptiness: 0x ) # Nitric Oxide (NO): A gas that diffuses freely.
- Nox ( full: 100x, active: 20x, empty: 0x ) # Nitric Oxide (NO): A gas that diffuses freely.
- Ecb ( fullness: 100x, active: 20x, emptiness: 0x ) # Endocannabinoids (e.g., 2-AG)
- Ecb ( full: 100x, active: 20x, empty: 0x ) # Endocannabinoids (e.g., 2-AG)
tub_intricated:
- Nt ( contained_by: BEH-SYN )
- bAp ( contained_by: BEH-SOMA )
```
### Context
### ms: behaviors POST
#### CheckConditions:Context
Contestualizziamo in maniera Fixed ogni mezzo secondo?
```Gen
context: captureNt
contained_by: BEH-POST
context: CheckConditions
contained_by: BEH-POST
in_context: Fixed
rf: ( active: 10x )
in_context: Fixed
rf: ( active: 600x )
condition: (Nt full) AND NOT (bAp)
out_context: NtCaptured
condition: NOT (RP empty) AND NOT (RRP full)
out_context: RPShuttle
condition: NOT (CaTrace empty)
out_context: CaTracesNotEmpty
condition: NOT (eCB empy)
out_context: eCBNotEmpty
```
### Episode
#### :Episode
## BEH-POST-AMPA: Container
@@ -114,7 +124,13 @@ container: BEH-POST-AMPA
- bAp ( contained_by: BEH-SOMA )
```
### AmpaOpen: Episode
### sec: behaviors POST
### min: behaviors POST
### ms: behaviors AMPA
#### AmpaOpen: Episode
- Timing: < 1 ms
- InContext: Glutamate > FULLNESS
@@ -134,7 +150,7 @@ episode: AmpaOpen
trace: None # Se Ca+FULLNESS, lascio tracce di overflow per modulazione DOWN, da capire UP
```
### Depolarization by bAP
#### Depolarization by bAP
Da ricordare i Dendritic VCGG che si aprono facendo entrare Ca2+ all'arrivo di bAP. In teoria abbiamo 3 Ion Channel.
@@ -142,23 +158,23 @@ Da ricordare i Dendritic VCGG che si aprono facendo entrare Ca2+ all'arrivo di b
- InContext: bAP backpropagating action potential
- Consequence: addition to local depolarization (EPSP) from bAP
### Mg²⁺ NMDA unblock
#### Mg²⁺ NMDA unblock
- Timing: > 1 ms
- InContext: local depolarization (EPSP) > FULLNESS (requires depolarization > -40mV)
- OutContext: NMDA Mg²⁺ unblock
### Full Ca+ influx by NDMA
#### Full Ca+ influx by NDMA
- Timing: < 1 ms
- InContext: NMDA Mg²⁺ unblock
- Consequence: Na⁺ influx
- Consequence: addition to local depolarization (EPSP) from Full NMDA activation
### Upregulation: Observable
#### Upregulation: Observable
- Upregulation: Depolarization relieves NMDA Mg²⁺ block → Ca²⁺ influx amplification
### Downregulation: Observable
#### Downregulation: Observable
- Downregulation: AMPA desensitization acts as low-pass filter
neuron/appunti/2026-03-30-behavior-POST.md
+276 -68
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@@ -1,91 +1,299 @@
## BEH-POST-UNIFIED: The Integrated Postsynaptic Model
---
### **ms: behaviors (Fast Kinetics & Gate Logic)**
**What needs adjustment**
#### **Voltage-Context: Episode**
`Voltage-Context` is labelled as an Episode but contains sub-episodes — it should be a Context. Same for `NMDA-Coincidence` and `Ca-Dynamics & ATP-Drain`. You have used Episode and Context interchangeably in a few places; the distinction matters for the specification: a Context sets the conditions, an Episode is a named outcome within those conditions.
*Determines the total depolarization level (Vpost​) available to clear the NMDA Mg-block.*
`Vpost_Maximum` lists `V_bAP full OR (g_AMPA full AND V_bAP medium)` — this is correct Boolean logic and should be kept exactly as written. It captures the two ways the postsynapse can reach maximum depolarisation: the bAP alone if it is strong enough, or AMPA plus a partial bAP together.
- **Vpost_Maximum: Episode**
- — `V_bAP` full OR (`g_AMPA` full AND `V_bAP` medium)
- — Result: Vpost​ is high enough for complete Mg-block removal.
- **Vpost_Attenuated: Episode**
- — `g_AMPA` medium AND `V_bAP` empty/low
- — Result: Vpost​ is sub-threshold; Mg-block partially remains.
- **Vpost_Passive: Episode**
- — `g_AMPA` empty AND `V_bAP` empty
- — Result: Vpost​ at rest; Mg-block fully intact.
`AMPA_Population_Increase` correctly gates on `ATP_level_post NOT empty` — LTP requires ATP for CaMKII phosphorylation and receptor trafficking. LTD does not have the same gate in your spec. This is slightly asymmetric — LTD (receptor internalisation via endocytosis) is also ATP-dependent, though less so than LTP. Worth noting.
#### **NMDA-Coincidence: Context**
`Plasticity_LTD` covers `Ca_post_history medium` but does not cover the case where `Ca_post_history` is empty — a completely silent synapse also weakens over time (homeostatic depression). This can be left as a gap or named explicitly.
- **NMDA_Open: Episode**
- — `NT_cleft` full AND `V_post` maximum
- — Result: Capost​ surges; triggers high ATP demand for clearance.
- **NMDA_LogicBlocked: Episode**
- — `NT_cleft` full BUT `V_post` attenuated/passive
- — Result: Mg-block prevents Ca2+ influx despite NT presence.
- **NMDA_LigandBlocked: Episode**
- — `V_post` maximum BUT `NT_cleft` empty
- — Result: No glutamate to open the channel; Ca2+ entry zero.
`Astrocyte_Supply_Crises` has a typo (Crises → Crisis).
#### **Ca-Dynamics & ATP-Drain: Context**
- **Clearance_Optimal: Episode**
- — `ATP_level_post` full
- — Result: Pumps flush Ca2+ rapidly; `ATP_demand_post` increases.
- **Clearance_Failing: Episode**
- — `ATP_level_post` low/empty
- — Result: Ca2+ remains elevated (residual floor); creates "False Trigger" conditions.
The bAP needs new parameters and state variables in the model before it can be expressed as a fully implemented behavior. I will add those inline as `[GAP — to implement]` markers so the spec is complete even where the code is not yet written.
---
### **sec: behaviors (Signal Integration & Fate)**
**Final restructured specification**
#### **Synaptic-Weight-Decision (Plasticity)**
```
BEH-POST-UNIFIED: The Integrated Postsynaptic Model
=====================================================
Three loops, three timescales, one shared astrocyte supply.
Each loop feeds the next: V_post enables NMDA, NMDA drives
Ca_post, Ca_post determines plasticity and eCB, ATP sustains
all three. Failure of ATP does not silence the postsynapse
(unlike the presynapse) — it corrupts it, triggering false
retrograde signals and risking excitotoxic Ca2+ accumulation.
- **Plasticity_LTP: Episode**
- — `Ca_post_history` (recent 2s) is **Full**
- — Result: High-frequency/High-amplitude coincidence detected; tags synapse for AMPA increase.
- **Plasticity_LTD: Episode**
- — `Ca_post_history` is **Medium**
- — Result: Low-frequency or poorly timed coincidence; tags synapse for AMPA removal.
Variables:
V_bAP — back-propagating AP amplitude (0→1)
[GAP — requires bAP_train input,
analogous to presynaptic spike_train]
g_AMPA — AMPA receptor conductance (= receptor_conductance)
V_post — total postsynaptic membrane potential (0→1)
NT_cleft — glutamate in cleft (from presynapse)
Desensitization — fraction of AMPA receptors desensitized (0→1)
Ca_post — free Ca2+ in postsynaptic spine (0→...)
Ca_post_history — 2 s rolling mean of Ca_post
ATP_level_post — normalised postsynaptic ATP (0→1)
ATP_demand_post — accumulated ATP cost since last metabolic cycle
g_AMPA_baseline — long-term AMPA receptor density set by plasticity
[GAP — not yet in model; LTP/LTD would write this]
eCB_level — endocannabinoid retrograde signal (0→1)
written here, read by presynapse Loop 1
#### **Retrograde-Feedback (eCB)**
━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━
ms: behaviors — Fast Kinetics and Gate Logic
━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━
- **eCB_Synthesis_Active: Episode**
- — `Ca_post_history` > threshold (0.7)
- — **Logic A (Protection):** Response to genuine NMDA over-activity.
- — **Logic B (Error):** Response to pump failure (Ca2+ accumulation due to low ATP).
- **eCB_Synthesis_Idle: Episode**
- — `Ca_post_history` < threshold; eCB level decays.
Voltage-Context: Context
Determines the total depolarisation (V_post) available to
lift the NMDA Mg block. Two independent sources contribute:
AMPA-driven local depolarisation (g_AMPA) and the somatic
back-propagating AP (V_bAP). Either alone can partially
depolarise; both together reach maximum.
---
Vpost_Maximum: Episode
— V_bAP full OR
— g_AMPA full AND V_bAP medium
— Result: V_post high enough for complete Mg block removal.
NMDA gate can open fully.
Both ATP costs charged at maximum rate.
### **min: behaviors (Bioenergetics & Structural Change)**
Vpost_Attenuated: Episode
— g_AMPA medium AND V_bAP empty/low OR
— g_AMPA low AND V_bAP medium
— Result: V_post sub-threshold.
Mg block partially remains.
NMDA gate opens partially or not at all.
This is the most common state during low-rate firing
without a coincident bAP.
#### **Metabolic-Recovery**
Vpost_Passive: Episode
— g_AMPA empty AND V_bAP empty
— Result: V_post at rest.
Mg block fully intact.
No Ca_post entry possible.
Na/K-ATPase cost minimal.
- **Astrocyte_Supply_Active: Episode**
- — `Glucose_level` full; `ATP_level_post` is replenished.
- **Astrocyte_Supply_Crises: Episode**
- — `Glucose_level` low; `ATP_level_post` remains empty; Ca2+ pumps fail.
Desensitization-Context: Context
Modulates g_AMPA independently of NT_cleft.
Sustained NT exposure drives receptors into a closed state
that persists even when NT remains present.
#### **Structural-Update**
DesensitizationRising: Episode
— NT_cleft sustained high (multiple consecutive ms)
— Desensitization rises each ms
— g_AMPA effectively reduced despite NT presence
— attenuates Vpost_Maximum toward Vpost_Attenuated
- **AMPA_Population_Increase: Episode**
- — Triggered by `Plasticity_LTP` AND `ATP_level_post` NOT empty.
- — Result: `g_AMPA_baseline` shifts higher for next cycle.
- **AMPA_Population_Decrease: Episode**
- — Triggered by `Plasticity_LTD`.
- — Result: `g_AMPA_baseline` shifts lower.
DesensitizationRecovering: Episode
— NT_cleft low or empty
— Desensitization decays with tau_desensitization = 500 ms
— g_AMPA ceiling restored gradually
---
NMDA-Coincidence: Context
Strict AND gate: both NT (ligand) and V_post (voltage) must
be simultaneously non-zero for Ca_post to rise.
Unlike presynaptic VGCCs which open with any spike, NMDA
requires coincidence. This makes Ca_post a detector of
coordinated pre+post activity, not just input rate.
### **Key Structural Observations**
NMDA_Open: Episode
— NT_cleft full AND V_post maximum (Vpost_Maximum active)
— Mg block fully lifted
— Ca_post surges — LTP territory
— ATP_demand_post rises sharply (PMCA must clear Ca_post)
— if sustained → Ca_post_history crosses eCB threshold
1. **The Vpost Priority:** Vpost​ is now the "sum" of local input and global feedback. This allows the model to simulate both **Hebbian learning** (needs a bAP) and **Homeostatic Scaling** (where enough local AMPA activity can eventually trigger the NMDA gate alone).
2. **The ATP Bottleneck for LTP:** Note that in the `min` behaviors, I’ve added a constraint: LTP requires `ATP_level_post` to be healthy. Moving receptors into the membrane is a physical "work" process. If the synapse is energy-starved, it might "decide" to do LTP in the `sec` loop but fail to **execute** it in the `min` loop.
3. **The Coincidence Hierarchy:**
- **LTP** = Perfect Timing (NT + bAP).
- **LTD** = Misaligned Timing (Low NT or Out-of-sync bAP).
- **False eCB** = Metabolic Failure (No NT, No bAP, just Low ATP).
NMDA_LogicBlocked: Episode
— NT_cleft full BUT V_post attenuated or passive
— Mg block partially or fully intact
— Ca_post does not rise despite NT presence
— Result: presynapse fired but postsynapse was not ready
No plasticity signal generated
This is the mechanism for input selectivity:
only synapses active during postsynaptic firing
produce a Ca_post signal
NMDA_LigandBlocked: Episode
— V_post maximum BUT NT_cleft empty
— No glutamate to open the channel
— Ca_post entry zero despite full depolarisation
— Result: bAP arrived but presynapse was silent
Again no plasticity signal
The AND logic enforces true coincidence
Ca-Dynamics-Context: Context
Ca_post clearance rate depends entirely on ATP_level_post.
This is the bridge from the ATP loop into the Ca2+ loop.
When ATP fails, Ca_post clearance fails, and the Ca2+ loop
becomes corrupted — Ca_post reflects pump state rather
than genuine coincidence events.
Clearance_Optimal: Episode
— ATP_level_post full → pump_scale_post near 1
— PMCA (ATP-gated) + NCX (always on) both clearing
— Ca_post returns to baseline between events
— Each coincidence event is temporally isolated
— ATP_demand_post increases proportionally to Ca_post load
Clearance_Reduced: Episode
— ATP_level_post medium → pump_scale_post reduced
— Ca_post clears more slowly
— Residual elevation begins accumulating between events
— Ca_post_history starts drifting upward
— eCB threshold may be approached during heavy firing
Clearance_Failing: Episode
— ATP_level_post low or empty → pump_scale_post near 0
— Only NCX clearing (floor, not rescue)
— Ca_post accumulates regardless of coincidence activity
— False Trigger conditions: Ca_post_history crosses eCB
threshold without genuine NMDA overactivity
— Excitotoxicity risk if Ca_post elevation is sustained
━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━
sec: behaviors — Signal Integration and Fate
━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━
Synaptic-Weight-Decision: Context
Ca_post_history (2 s rolling mean of Ca_post) determines
the plasticity signal. The threshold logic is graded:
the same variable produces opposite outcomes depending
on whether it is above or below the LTP/LTD boundary.
ATP_level_post gates LTP expression but not LTD —
strengthening requires energy, weakening does not.
Plasticity_LTP: Episode
— Ca_post_history full (above Ca_post_LTP threshold)
— High-frequency or high-amplitude coincidence detected
— Tags synapse for AMPA receptor insertion
— Requires ATP_level_post NOT empty for expression
(CaMKII phosphorylation and receptor trafficking are
ATP-dependent — energy failure blocks LTP even if
the Ca_post signal is correct)
— [GAP] LTP expression writes g_AMPA_baseline upward
in the minutes loop
Plasticity_Boundary: Episode
— Ca_post_history medium
— Poorly timed or low-frequency coincidence
— Neither LTP nor LTD threshold crossed
— Synapse weight unchanged this cycle
Plasticity_LTD: Episode
— Ca_post_history low but non-zero
— Weak or mistimed coincidence — presynapse fired
but postsynapse was not sufficiently depolarised
— Tags synapse for AMPA receptor removal
— Less ATP-dependent than LTP; can proceed under
mild energy stress
— [GAP] LTD expression writes g_AMPA_baseline downward
in the minutes loop
Plasticity_Silent: Episode
— Ca_post_history empty (prolonged absence of activity)
— Homeostatic depression: unused synapses weaken
— [GAP] not yet modelled; would require Ca_post_trace
integration over hours
Retrograde-Feedback: Context
eCB synthesis is triggered by Ca_post_history, not V_post.
It is Ca2+ in the spine — not voltage — that activates the
enzymes (DAGL, PLC) that produce endocannabinoids.
The model cannot distinguish internally between the two
causes of elevated Ca_post_history (genuine vs pump failure)
but the consequences differ: one is communication,
the other is survival.
eCB_Synthesis_Active: Episode
— Ca_post_history > eCB_threshold (0.7)
— Logic A (Genuine Protection):
Cause : sustained NMDA_Open events — real overactivity
Effect : appropriate retrograde stop signal
Outcome : presynapse reduces NT → NT_cleft falls →
NMDA closes → Ca_post load drops →
Ca_post_history falls → eCB synthesis subsides
Loop closes correctly
— Logic B (False Trigger — Excitotoxic Protection):
Cause : Clearance_Failing — Ca_post elevated by
pump failure, not genuine coincidence
Effect : presynapse silenced without real overactivity
Outcome : NT_cleft falls → NMDA closes → Ca_post
load drops → ATP_demand_post falls →
ATP_level_post recovers → pumps restart →
Ca_post clears → Ca_post_history falls →
eCB synthesis subsides
Desperate survival loop — buys time for
metabolic recovery
eCB_Synthesis_Idle: Episode
— Ca_post_history < eCB_threshold
— eCB_level decays with tau_eCB_decay = 10000 ms
— Presynaptic suppression lifts gradually
— 10 s decay means suppression outlasts the trigger —
prevents immediate re-engagement before Ca_post
has stabilised
━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━
min: behaviors — Bioenergetics and Structural Change
━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━━
Metabolic-Recovery: Context
ATP_level_post is computed from Glucose_level (shared
supply) minus ATP_demand_post (postsynaptic-specific cost).
The shared supply creates the coupling: both pre and post
deplete the same astrocyte glucose budget simultaneously.
Presynaptic silence is therefore metabolically beneficial
to the postsynapse — less NT means less NMDA activation
means less Ca_post means less PMCA cost.
Astrocyte_Supply_Active: Episode
— Glucose_level full
— ATP_demand_post within supply capacity
— ATP_level_post replenished each cycle
— All three loops operating normally
Astrocyte_Supply_Stressed: Episode
— Glucose_level medium OR ATP_demand_post elevated
— ATP_level_post partially reduced
— Clearance_Reduced begins
— Plasticity_LTP at risk (ATP dependency)
Astrocyte_Supply_Crisis: Episode
— Glucose_level low OR ATP_demand_post chronically high
— ATP_level_post near empty
— Clearance_Failing — only NCX clearing Ca_post
— False Trigger likely → eCB_Synthesis_Active (Logic B)
— Presynaptic silence indirectly reduces postsynaptic
ATP demand — the coupled protection mechanism
Structural-Update: Context
Long-term changes to AMPA receptor density.
These are the physical substrate of learning and memory.
[GAP] g_AMPA_baseline not yet implemented in the model.
Would be written in Loop 3 and read by Loop 1 as the
ceiling of receptor_conductance.
AMPA_Population_Increase: Episode
— Triggered by Plasticity_LTP
— AND ATP_level_post NOT empty
— g_AMPA_baseline shifts higher for next cycle
— More AMPA receptors → stronger Vpost_Maximum
— Easier to reach NMDA_Open in future events
— Positive feedback: LTP makes future LTP more likely
AMPA_Population_Decrease: Episode
— Triggered by Plasticity_LTD or Plasticity_Silent
— g_AMPA_baseline shifts lower
— Fewer AMPA receptors → Vpost_Attenuated more common
— Harder to reach NMDA_Open → weakens synapse further
— Negative feedback: LTD stabilises by reducing
future coincidence probability
```