281 lines
24 KiB
HTML
281 lines
24 KiB
HTML
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<style>
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.root { padding: 1rem 0; font-family: var(--font-mono); font-size: 13px; color: var(--color-text-primary); line-height: 1.75; }
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.section { margin-bottom: 2rem; }
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.section-header { font-size: 11px; font-weight: 500; letter-spacing: 0.08em; text-transform: uppercase; color: var(--color-text-tertiary); border-bottom: 1px solid var(--color-border-tertiary); padding-bottom: 4px; margin-bottom: 0.75rem; }
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.block { border-left: 2px solid var(--color-border-secondary); padding-left: 1rem; margin-bottom: 1.25rem; }
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.kw { color: var(--color-text-info); }
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.fn { color: var(--color-text-warning); }
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.cm { color: var(--color-text-tertiary); font-style: italic; }
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.pre { color: #c084fc; }
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.pst { color: var(--color-text-success); }
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.ast { color: #f0c070; }
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.sig { color: var(--color-text-info); }
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.num { color: var(--color-text-danger); }
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.i1 { padding-left: 1.5rem; }
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.i2 { padding-left: 3rem; }
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.i3 { padding-left: 4.5rem; }
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.sep { border: none; border-top: 1px solid var(--color-border-tertiary); margin: 2rem 0; }
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.legend { display: flex; gap: 1.25rem; flex-wrap: wrap; margin-bottom: 1.25rem; font-size: 11px; padding: 8px 12px; background: var(--color-background-secondary); border-radius: 6px; }
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.leg { display: flex; align-items: center; gap: 5px; }
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.dot { width: 8px; height: 8px; border-radius: 50%; flex-shrink: 0; }
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</style>
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<div class="root">
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<div class="legend">
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<div class="leg"><div class="dot" style="background:var(--color-text-success)"></div><span>postsynapse</span></div>
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<div class="leg"><div class="dot" style="background:#c084fc"></div><span>presynapse</span></div>
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<div class="leg"><div class="dot" style="background:#f0c070"></div><span>astrocyte</span></div>
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<div class="leg"><div class="dot" style="background:var(--color-text-info)"></div><span>signal / neuromodulator</span></div>
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</div>
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<div class="section">
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<div class="section-header">global state variables</div>
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<div class="block">
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<div><span class="cm">// ── Fast (ms–s): wave propagation ─────────────────────────────</span></div>
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<div><span class="cm">// Presynapse</span></div>
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<div><span class="pre">pre_Ca_residual</span> <span class="cm">// leftover Ca²⁺ between spikes — short-term trace</span></div>
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<div><span class="pre">vesicle_release_prob</span> <span class="cm">// P(0.1–1.0) per docking slot</span></div>
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<div><span class="pre">RRP_pool</span> <span class="cm">// readily-releasable vesicle pool</span></div>
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<div><span class="pre">reserve_pool</span> <span class="cm">// chained vesicles in deep storage</span></div>
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<br>
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<div><span class="cm">// Postsynapse</span></div>
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<div><span class="pst">membrane_potential</span> <span class="cm">// Vm — depolarization state</span></div>
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<div><span class="pst">NMDA_Mg_block</span> <span class="cm">// bool — mechanical clamp on/off</span></div>
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<div><span class="pst">post_Ca_amplitude</span> <span class="cm">// peak [Ca²⁺] rise in spine</span></div>
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<div><span class="pst">post_Ca_rise_speed</span> <span class="cm">// d(Ca)/dt — fast=LTP signal, slow=LTD signal</span></div>
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<br>
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<div><span class="cm">// Astrocyte</span></div>
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<div><span class="ast">glutamate_cleft</span> <span class="cm">// [glu] in synaptic cleft</span></div>
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<div><span class="ast">glutamate_spillover</span> <span class="cm">// extrasynaptic [glu] — saturates mGluRs</span></div>
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<div><span class="ast">astro_Ca_local</span> <span class="cm">// IP3-triggered local rise near synapse</span></div>
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<div><span class="ast">astro_Ca_global</span> <span class="cm">// soma-wide wave — network overload flag</span></div>
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<div><span class="ast">D_serine_release</span> <span class="cm">// gliotransmitter — NMDA co-agonist pulse</span></div>
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<div><span class="ast">lactate_output</span> <span class="cm">// fuel export rate to pre and post</span></div>
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<br>
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<div><span class="cm">// ── Intermediate (s–min): temporary tuning ────────────────────</span></div>
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<div><span class="sig">mGluR2_3_activation</span> <span class="cm">// presynaptic Gi — autoinhibitory brake</span></div>
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<div><span class="sig">mGluR5_activation</span> <span class="cm">// astrocytic Gq — IP3→Ca²⁺→D-serine cascade</span></div>
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<div><span class="sig">cAMP_level</span> <span class="cm">// set by dopamine/NE via Gs → adenylyl cyclase</span></div>
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<div><span class="sig">PKA_activity</span> <span class="cm">// downstream of cAMP</span></div>
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<div><span class="sig">GluA1_Ser845_primed</span> <span class="cm">// bool — AMPA insertion threshold lowered by PKA</span></div>
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<div><span class="sig">DARPP32_phospho</span> <span class="cm">// bool — PP1 (LTD phosphatase) silenced by PKA</span></div>
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<div><span class="sig">CREB_active</span> <span class="cm">// bool — structural gene expression enabled</span></div>
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<br>
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<div><span class="cm">// ── Slow (h–weeks): structural architecture ───────────────────</span></div>
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<div><span class="pst">AMPA_count</span> <span class="cm">// surface receptors — postsynaptic sensitivity</span></div>
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<div><span class="pst">spine_volume</span> <span class="cm">// physical size of dendritic spine</span></div>
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<div><span class="pre">active_zone_size</span> <span class="cm">// docking slot count</span></div>
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<div><span class="pre">RRP_pool_capacity</span> <span class="cm">// max readily-releasable pool</span></div>
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<div><span class="pre">VGCC_clustering</span> <span class="cm">// Ca²⁺ channels beneath active zone</span></div>
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<div><span class="ast">perisynaptic_distance</span> <span class="cm">// how close astrocyte walls are to synapse</span></div>
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<div><span class="ast">ECM_integrity</span> <span class="cm">// extracellular matrix density</span></div>
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<div><span class="ast">D_serine_tonic_level</span> <span class="cm">// baseline co-agonist supply (sustained)</span></div>
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<div><span class="ast">glutamate_clearance_rate</span> <span class="cm">// EAAT transporter density</span></div>
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</div>
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</div>
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<hr class="sep">
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<div class="section">
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<div class="section-header">fast time scale — wave propagation (ms → s)</div>
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<div class="block">
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<div><span class="kw">function</span> <span class="fn">fire_action_potential</span>(input_freq):</div>
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<br>
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<div class="i1"><span class="cm">// Presynapse: launch wavefront</span></div>
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<div class="i1"><span class="pre">pre_Ca_residual</span> += spike_influx(input_freq)</div>
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<div class="i1"><span class="pre">pre_Ca_residual</span> *= decay(τ ≈ <span class="num">100ms</span>) <span class="cm">// fades unless spikes keep arriving</span></div>
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<div class="i1"><span class="pre">vesicle_release_prob</span> *= facilitation(<span class="pre">pre_Ca_residual</span>)</div>
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<div class="i1">released_vesicles = binomial(<span class="pre">RRP_pool</span>, <span class="pre">vesicle_release_prob</span>)</div>
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<div class="i1"><span class="ast">glutamate_cleft</span> = released_vesicles × quantal_content</div>
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<div class="i1"><span class="pre">RRP_pool</span> -= released_vesicles</div>
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<br>
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<div class="i1"><span class="cm">// Astrocyte: overflow sensing and co-agonist release</span></div>
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<div class="i1"><span class="ast">glutamate_spillover</span> = extrasynaptic_diffusion(<span class="ast">glutamate_cleft</span>)</div>
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<div class="i1"><span class="kw">if</span> <span class="ast">glutamate_spillover</span> > spillover_threshold:</div>
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<div class="i2"><span class="sig">mGluR5_activation</span> = <span class="kw">True</span> <span class="cm">// Gq arm → IP3 → Ca²⁺ → D-serine</span></div>
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<div class="i2"><span class="ast">astro_Ca_local</span> += IP3_cascade(PLC)</div>
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<div class="i2"><span class="ast">D_serine_release</span> += proportional_to(<span class="ast">astro_Ca_local</span>)</div>
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<div class="i2"><span class="sig">mGluR2_3_activation</span> = <span class="kw">True</span> <span class="cm">// Gi arm → brake presynapse</span></div>
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<div class="i2"><span class="sig">cAMP_level</span> -= Gi_inhibition(adenylyl_cyclase)</div>
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<div class="i2"><span class="pre">vesicle_release_prob</span> -= VGCC_suppression() <span class="cm">// autoinhibitory brake</span></div>
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<br>
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<div class="i1"><span class="cm">// Astrocyte: check for network overload</span></div>
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<div class="i1"><span class="ast">astro_Ca_global</span> = soma_wave(<span class="ast">astro_Ca_local</span> > OVERLOAD_threshold)</div>
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<div class="i1"><span class="kw">if</span> <span class="ast">astro_Ca_global</span>: <span class="fn">trigger</span>(shockwave_lockdown)</div>
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<br>
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<div class="i1"><span class="cm">// Postsynapse: wavefront strikes resonator</span></div>
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<div class="i1">AMPA_current = <span class="ast">glutamate_cleft</span> × <span class="pst">AMPA_count</span></div>
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<div class="i1"><span class="pst">membrane_potential</span> += AMPA_current</div>
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<br>
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<div class="i1"><span class="cm">// NMDA gate: coincidence check</span></div>
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<div class="i1"><span class="kw">if</span> <span class="pst">membrane_potential</span> > <span class="num">-40mV</span> <span class="kw">and</span> <span class="ast">D_serine_release</span> > threshold:</div>
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<div class="i2"><span class="pst">NMDA_Mg_block</span> = <span class="kw">False</span> <span class="cm">// Mg²⁺ ejected</span></div>
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<div class="i2"><span class="pst">post_Ca_amplitude</span> += NMDA_influx(<span class="ast">glutamate_cleft</span>)</div>
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<div class="i2"><span class="pst">post_Ca_rise_speed</span> = d(<span class="pst">post_Ca_amplitude</span>) / dt</div>
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<br>
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<div class="i1"><span class="cm">// Astrocyte: vacuum trailing echoes + fuel pipeline</span></div>
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<div class="i1"><span class="ast">glutamate_cleft</span> -= <span class="ast">glutamate_clearance_rate</span> × Δt</div>
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<div class="i1"><span class="ast">lactate_output</span> += glycolysis_rate(<span class="ast">glutamate_clearance_rate</span>)</div>
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<div class="i1"><span class="pst">membrane_potential</span> restored by NaK_ATPase(<span class="ast">lactate_output</span>)</div>
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<div class="i1"><span class="pre">RRP_pool</span> refilled by VATPase(<span class="ast">lactate_output</span>)</div>
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</div>
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</div>
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<hr class="sep">
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<div class="section">
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<div class="section-header">intermediate time scale — temporary tuning (s → min)</div>
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<div class="block">
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<div><span class="kw">function</span> <span class="fn">short_term_plasticity</span>(input_freq, duration):</div>
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<br>
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<div class="i1"><span class="cm">// Presynapse: facilitate or depress based on Ca²⁺ history</span></div>
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<div class="i1"><span class="kw">if</span> input_freq > <span class="num">20Hz</span>:</div>
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<div class="i2"><span class="pre">vesicle_release_prob</span> *= <span class="num">1.3</span> <span class="cm">// residual Ca²⁺ primes launchpad</span></div>
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<div class="i2"><span class="fn">mobilize</span>(<span class="pre">reserve_pool</span> → <span class="pre">RRP_pool</span>) <span class="cm">// break storage chains</span></div>
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<div class="i1"><span class="kw">elif</span> input_freq < <span class="num">5Hz</span>:</div>
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<div class="i2"><span class="pre">vesicle_release_prob</span> *= <span class="num">0.7</span> <span class="cm">// RRP depleted faster than refill</span></div>
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<br>
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<div class="i1"><span class="cm">// Postsynapse: NMDA gate primed if frequency sustained</span></div>
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<div class="i1"><span class="kw">if</span> input_freq >= <span class="num">50Hz</span> <span class="kw">and</span> duration > <span class="num">1s</span>:</div>
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<div class="i2"><span class="pst">NMDA_Mg_block</span> = <span class="kw">False</span> <span class="cm">// sustained depolarization</span></div>
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<div class="i2"><span class="pst">post_Ca_amplitude</span> accumulates <span class="cm">// early-LTP signal rises</span></div>
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<br>
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<div class="i1"><span class="cm">// Astrocyte: sustained volume → escalate co-agonist</span></div>
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<div class="i1"><span class="kw">if</span> <span class="ast">astro_Ca_local</span> > local_threshold:</div>
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<div class="i2"><span class="ast">D_serine_release</span> += gliotransmitter_pulse() <span class="cm">// widens NMDA window</span></div>
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<br>
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<div class="i1"><span class="cm">// Neuromodulators: set context gate via Gs protein</span></div>
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<div class="i1"><span class="kw">if</span> dopamine_level > D1_threshold <span class="kw">or</span> NE_level > β_threshold:</div>
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<div class="i2"><span class="sig">cAMP_level</span> += Gs_activation(adenylyl_cyclase)</div>
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<div class="i2"><span class="sig">PKA_activity</span> = proportional_to(<span class="sig">cAMP_level</span>)</div>
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<div class="i2"><span class="fn">phosphorylate</span>(GluA1, site=Ser845)</div>
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<div class="i2"><span class="sig">GluA1_Ser845_primed</span> = <span class="kw">True</span> <span class="cm">// lowers CaMKII threshold</span></div>
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<div class="i2"><span class="fn">phosphorylate</span>(DARPP32)</div>
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<div class="i2"><span class="sig">DARPP32_phospho</span> = <span class="kw">True</span> <span class="cm">// silences PP1 — blocks LTD</span></div>
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<div class="i2"><span class="fn">translocate</span>(PKA → nucleus) → <span class="fn">phosphorylate</span>(CREB)</div>
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<div class="i2"><span class="sig">CREB_active</span> = <span class="kw">True</span> <span class="cm">// enables structural gene expression</span></div>
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<div class="i1"><span class="cm">// Acetylcholine: lower LTP threshold globally</span></div>
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<div class="i1">LTP_threshold *= (<span class="num">1</span> / (<span class="num">1</span> + ACh_level × mAChR_gain))</div>
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</div>
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</div>
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<hr class="sep">
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<div class="section">
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<div class="section-header">slow time scale — structural commit (h → weeks)</div>
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<div class="block">
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<div><span class="kw">function</span> <span class="fn">commit_to_structural_change</span>():</div>
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<br>
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<div class="i1"><span class="cm">// Hierarchical filter: three conditions must align</span></div>
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<div class="i1">event_detected = <span class="pst">post_Ca_amplitude</span> > Ca_HIGH <span class="cm">// layer 1: did something happen?</span></div>
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<div class="i1">overflow_sensed = <span class="sig">mGluR5_activation</span> == <span class="kw">True</span> <span class="cm">// layer 2: was it excessive?</span></div>
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<div class="i1">context_validated = <span class="sig">DARPP32_phospho</span> <span class="kw">and</span> <span class="sig">GluA1_Ser845_primed</span> <span class="cm">// layer 3: worth saving?</span></div>
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<br>
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<div class="i1"><span class="cm">// ── Branch 1: LTP — potentiation ──────────────────────────────</span></div>
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<div class="i1"><span class="kw">if</span> event_detected <span class="kw">and</span> overflow_sensed <span class="kw">and</span> context_validated:</div>
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<br>
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<div class="i2"><span class="cm">// Postsynapse: anchor receptors, enlarge spine</span></div>
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<div class="i2"><span class="fn">activate</span>(CaMKII)</div>
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<div class="i2"><span class="pst">AMPA_count</span> += receptor_insertion(CaMKII, <span class="sig">GluA1_Ser845_primed</span>)</div>
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<div class="i2"><span class="pst">spine_volume</span> *= <span class="num">1.5</span></div>
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<br>
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<div class="i2"><span class="cm">// Presynapse: expand launchpad, increase output reliability</span></div>
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<div class="i2"><span class="pre">active_zone_size</span> *= <span class="num">1.4</span> <span class="cm">// more docking slots</span></div>
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<div class="i2"><span class="pre">RRP_pool_capacity</span> += pool_expansion(<span class="pre">active_zone_size</span>)</div>
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<div class="i2"><span class="pre">VGCC_clustering</span> += cluster_beneath_AZ() <span class="cm">// tighter Ca²⁺ coupling</span></div>
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<div class="i2"><span class="pre">vesicle_release_prob</span> += <span class="num">0.1</span> <span class="cm">// driven by VGCC clustering</span></div>
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<br>
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<div class="i2"><span class="cm">// Astrocyte: seal and insulate the channel</span></div>
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<div class="i2"><span class="ast">perisynaptic_distance</span> -= process_retraction() <span class="cm">// walls move IN → tighter wrap</span></div>
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<div class="i2"><span class="ast">ECM_integrity</span> += secrete(Glypicans, Thrombospondins)</div>
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<div class="i2"><span class="ast">D_serine_tonic_level</span> += upregulate_synthesis() <span class="cm">// sustained NMDA priming</span></div>
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<div class="i2"><span class="ast">glutamate_clearance_rate</span> *= <span class="num">0.85</span> <span class="cm">// tighter wrap → slower diffusion away</span></div>
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<div class="i2"><span class="kw">return</span> <span class="num">"potentiated"</span></div>
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<br>
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<div class="i1"><span class="cm">// ── Branch 2: temporary only — Ca²⁺ rose, no save signal ─────</span></div>
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<div class="i1"><span class="kw">elif</span> event_detected <span class="kw">and</span> <span class="kw">not</span> context_validated:</div>
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<div class="i2"><span class="pst">AMPA_count</span> += transient_insertion() <span class="cm">// early-LTP only — reverses in minutes</span></div>
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<div class="i2"><span class="pre">vesicle_release_prob</span> += transient_facilitation()</div>
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<div class="i2"><span class="cm">// No astrocyte structural change</span></div>
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<div class="i2"><span class="kw">return</span> <span class="num">"temporary facilitation only"</span></div>
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<br>
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<div class="i1"><span class="cm">// ── Branch 3: LTD — active forgetting ─────────────────────────</span></div>
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<div class="i1"><span class="kw">elif</span> event_detected <span class="kw">and</span> <span class="kw">not</span> overflow_sensed <span class="kw">and</span> <span class="kw">not</span> context_validated:</div>
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<br>
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<div class="i2"><span class="cm">// Postsynapse: internalize receptors, shrink spine</span></div>
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<div class="i2"><span class="fn">activate</span>(PP1)</div>
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<div class="i2"><span class="pst">AMPA_count</span> -= receptor_internalization(PP1)</div>
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<div class="i2"><span class="pst">spine_volume</span> *= <span class="num">0.7</span></div>
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<br>
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<div class="i2"><span class="cm">// Presynapse: dismantle launchpad</span></div>
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<div class="i2"><span class="pre">active_zone_size</span> -= docking_slot_removal()</div>
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<div class="i2"><span class="pre">RRP_pool_capacity</span> -= pool_contraction()</div>
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<div class="i2"><span class="pre">VGCC_clustering</span> -= scatter_VGCCs() <span class="cm">// decouple Ca²⁺ from AZ</span></div>
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<div class="i2"><span class="pre">vesicle_release_prob</span> *= <span class="num">0.6</span></div>
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<br>
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<div class="i2"><span class="cm">// Astrocyte: dissolve matrix, pull away, cut support</span></div>
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<div class="i2"><span class="ast">ECM_integrity</span> -= secrete(MMPs) <span class="cm">// molecular scissors</span></div>
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<div class="i2"><span class="ast">D_serine_tonic_level</span> = <span class="num">0</span> <span class="cm">// co-agonist supply cut</span></div>
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<div class="i2"><span class="ast">perisynaptic_distance</span> += process_extension() <span class="cm">// walls move OUT → loose wrap</span></div>
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<div class="i2"><span class="ast">glutamate_clearance_rate</span> *= <span class="num">1.2</span> <span class="cm">// looser wrap → faster spillover</span></div>
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<div class="i2"><span class="kw">return</span> <span class="num">"depressed"</span></div>
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<br>
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<div class="i1"><span class="cm">// ── Branch 4: baseline ────────────────────────────────────────</span></div>
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<div class="i1"><span class="kw">else</span>:</div>
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<div class="i2"><span class="cm">// All structural variables unchanged — system holds current state</span></div>
|
||
<div class="i2"><span class="kw">return</span> <span class="num">"baseline — no change"</span></div>
|
||
</div>
|
||
</div>
|
||
|
||
<hr class="sep">
|
||
|
||
<div class="section">
|
||
<div class="section-header">special case — shockwave lockdown (>100Hz uncoordinated)</div>
|
||
<div class="block">
|
||
<div><span class="kw">function</span> <span class="fn">shockwave_lockdown</span>():</div>
|
||
<div class="i1"><span class="ast">astro_Ca_global</span> = GLOBAL_WAVE <span class="cm">// soma-level flood</span></div>
|
||
<div class="i1"><span class="fn">release</span>(GABA, ATP) <span class="cm">// gel floods postsynapse</span></div>
|
||
<div class="i1"><span class="pst">AMPA_count</span> -= mass_internalization()</div>
|
||
<div class="i1"><span class="pst">membrane_potential</span> = HYPERPOLARIZED</div>
|
||
<div class="i1"><span class="fn">cluster</span>(<span class="pre">VGCC</span> → beneath_active_zone) <span class="cm">// ensures signal survives chaos</span></div>
|
||
</div>
|
||
</div>
|
||
|
||
<hr class="sep">
|
||
|
||
<div class="section">
|
||
<div class="section-header">energy supply chain — metabolic gating (continuous)</div>
|
||
<div class="block">
|
||
<div><span class="kw">function</span> <span class="fn">metabolic_loop</span>(Δt):</div>
|
||
<div class="i1"><span class="cm">// Astrocyte: glucose → lactate pipeline</span></div>
|
||
<div class="i1">glucose_uptake = blood_capillary_supply()</div>
|
||
<div class="i1"><span class="ast">lactate_output</span> = glycolysis(glucose_uptake, <span class="ast">glutamate_clearance_rate</span>)</div>
|
||
<div class="i1"><span class="ast">lactate_output</span> *= load_factor(<span class="ast">glutamate_clearance_rate</span>)</div>
|
||
<br>
|
||
<div class="i1"><span class="cm">// Pre + post absorb lactate → power their pumps</span></div>
|
||
<div class="i1"><span class="pre">RRP_pool</span> refill rate ∝ VATPase(<span class="ast">lactate_output</span>)</div>
|
||
<div class="i1"><span class="pst">membrane_potential</span> reset ∝ NaK_ATPase(<span class="ast">lactate_output</span>)</div>
|
||
</div>
|
||
</div>
|
||
|
||
<hr class="sep">
|
||
|
||
<div class="section">
|
||
<div class="section-header">key asymmetry — perisynaptic distance is bidirectional</div>
|
||
<div class="block">
|
||
<div><span class="cm">// LTP: astrocyte moves IN → tighter diffusion barrier</span></div>
|
||
<div><span class="cm">// → glutamate_clearance_rate ↓ (signal contained, not diluted)</span></div>
|
||
<div><span class="cm">// → D_serine_tonic_level ↑ (NMDA gate chronically primed)</span></div>
|
||
<br>
|
||
<div><span class="cm">// LTD: astrocyte moves OUT → looser diffusion barrier</span></div>
|
||
<div><span class="cm">// → glutamate_clearance_rate ↑ (signal bleeds away faster)</span></div>
|
||
<div><span class="cm">// → D_serine_tonic_level = 0 (NMDA gate chronically starved)</span></div>
|
||
<br>
|
||
<div><span class="cm">// Result: astrocyte amplifies both directions simultaneously</span></div>
|
||
<div><span class="cm">// potentiation becomes self-reinforcing; depression becomes self-reinforcing</span></div>
|
||
</div>
|
||
</div>
|
||
|
||
</div>
|