Introduction: The State-Dependence Problem in Neural Circuit Function

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^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference} A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference} A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...
^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference} A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...
^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference} The phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference} The phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...
^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference} The phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference} The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference} The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}0 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}1. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}2 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}3. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}4 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}5. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}6 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}7. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}8 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}9. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}0 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}1. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}2 The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete ^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}3. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...
^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}4 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}5 and dLight variants for dopamine ^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}6 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}7 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}8 and dLight variants for dopamine ^{3Citationpaper:paper-e13b61a8ecb6A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}9 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}0 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}1 and dLight variants for dopamine ^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}2 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}3 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}4 and dLight variants for dopamine ^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}5 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}6 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}7 and dLight variants for dopamine ^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}8 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{4Citationpaper:paper-58026432307eThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}9 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}0 and dLight variants for dopamine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}1 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}2 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}3 and dLight variants for dopamine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}4 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}5 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}6 and dLight variants for dopamine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}7 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}8 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}9 and dLight variants for dopamine ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}0 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}1 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}2 and dLight variants for dopamine ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}3 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}4 The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}5 and dLight variants for dopamine ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}6 — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...
^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}7 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}8. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{5Citationpaper:paper-a99781d55b5dThe phenomenon has been recognized for decades in invertebrate preparations, where the same small circuit can generate entirely different motor patterns depending on which neuromodulators are present in the surrounding milieu. Yet the implications of state-dependence have been slow to permeate the mainstream of mammalian systems neuroscience. Much of the field operates on an implicit assumption that circuit function...content/01_introduction.md:line 7Open reference}9. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}0 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}1. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}2. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}3 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}4. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}5. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}6 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}7. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}8. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}9 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}0. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}1. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}2 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}3. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}4. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}5 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}6. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}7. Serotoni...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}8 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}9. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}0. Serotoni...
^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}1 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}2. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}3. Serotoni...
^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}4 The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field’s understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson’s disease and addiction ^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}5. Acetylcholine research has been shaped by its link to Alzheimer’s disease and cortical attention ^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}6. Serotoni...
^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}7 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}8. Serotonin acting th...
^{6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}9 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}0. Serotonin acting th...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}1 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}2. Serotonin acting th...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}3 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}4. Serotonin acting th...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}5 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}6. Serotonin acting th...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}7 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}8. Serotonin acting th...
^{1Citationpaper:paper-1d9a4f7fb30aA neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference, 6Citationpaper:paper-c033caf877e7The computational consequences of state-dependence are profound. If the same circuit implements different computations under different modulatory conditions, then no single-state characterization of that circuit is complete [Guarino2025, Rahman2025b]. The reward prediction error signals attributed to dopamine neurons in the ventral tegmental area represent a computation that depends not only on the synaptic inputs t...content/01_introduction.md:line 9Open reference}9 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}00. Serotonin acting th...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}01 These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}02. Serotonin acting th...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}03 The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different “function” depending on which brain region the reviewer chose to emphasize. Dopamine is a “reward signal” if you read the striatal literature, a “working memory modulator” if you read the prefrontal literature, and a “salience detector” if you read the amygdala literat...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}04 The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different “function” depending on which brain region the reviewer chose to emphasize. Dopamine is a “reward signal” if you read the striatal literature, a “working memory modulator” if you read the prefrontal literature, and a “salience detector” if you read the amygdala literat...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}05 The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different “function” depending on which brain region the reviewer chose to emphasize. Dopamine is a “reward signal” if you read the striatal literature, a “working memory modulator” if you read the prefrontal literature, and a “salience detector” if you read the amygdala literat...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}06 The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different “function” depending on which brain region the reviewer chose to emphasize. Dopamine is a “reward signal” if you read the striatal literature, a “working memory modulator” if you read the prefrontal literature, and a “salience detector” if you read the amygdala literat...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}07 This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}08. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}09. The prefrontal cortex r...
^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}10 This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}11. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons ^{2Citationpaper:paper-6275c3b6b837A neural circuit is not a fixed input–output device. The same population of neurons, receiving the same afferent drive, can produce qualitatively different patterns of activity — and qualitatively different behavioral outcomes — depending on the modulatory state in which it operates. A cortical column that amplifies sensory signals during focused attention may suppress those same signals during drowsiness. A striata...content/01_introduction.md:line 5Open reference}12. The prefrontal cortex r...
... 47 additional anchors in refs_json

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[Wei2025b] paper:paper-6a24ab8dc72f “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[Labouesse2024] paper:paper-8eda8b8cc145 “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
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[Li2026c] paper:paper-fe4ebb70a6fe “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[Roth2018] paper:paper-b35554586e4b “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[Berthoux2025] paper:paper-2de7151e0d43 “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[Shanazz2025] paper:paper-d24cb9669281 “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[SanchezLeon2023] paper:paper-fced3ca3101b “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[Hickman2025] paper:paper-ce2af2624305 “The tools available to investigate state-dependence have undergone a revolution in the past decade. Genetically encoded neurotransmitter sensors — including the GRAB family for acetylcholine, dopamine, serotonin, and norepinephrine [Xie2024, Wei2025b] and dLight variants for dopamine [Labouesse2024, Tian2025] — now permit sub-second monitoring of modulatory dynamics in behaving animals, replacing the minutes-resolut...”
[Eisenstein2024] paper:paper-407d56ad8347 “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Stocchi2025] paper:paper-cbe116ba6b00 “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Hampel2018] paper:86682f3e-dd6a-4ba9-a2ea-8567b188ce7b “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Hampel2021] paper:13586346-e375-4206-9d52-342059b7134d “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Selkoe2016] paper:c7654776-8739-46c5-ad06-8d8edb3b7959 “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Moncrieff2023] paper:paper-cd570706bad6 “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Otte2016] paper:paper-7b3b5741091e “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[TodorovicVukotic2026] paper:paper-c853f88667e8 “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Caestecker2025a] paper:paper-02f6419b74fa “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[Waterhouse2022] paper:paper-5b5b7e051703 “The study of neuromodulation has been shaped by a set of historical accidents that continue to distort the field's understanding. Dopamine research has been dominated by the striatum, owing to its relevance for Parkinson's disease and addiction [Eisenstein2024, Stocchi2025]. Acetylcholine research has been shaped by its link to Alzheimer's disease and cortical attention [Hampel2018, Hampel2021, Selkoe2016]. Serotoni...”
[DEsposito2015] paper:paper-fa7ca0dd13a2 “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Shao2025] paper:paper-f2caf483b95b “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Hamada2024] paper:paper-62d25a818bc9 “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Mackowiak2023] paper:paper-f654c44bb6c5 “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Nieuwenhuis2005] paper:paper-397bd7afbb90 “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Dolan2002] paper:paper-5b095bc2e4f1 “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Kim2025b] paper:paper-17f807840d5a “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[Kaganovski2025] paper:paper-e2b2271c3841 “These associations are not wrong, but they are dangerously incomplete. Each neuromodulatory system acts across virtually the entire brain, and the effects of a given modulator can differ qualitatively between target regions. Dopamine in the prefrontal cortex follows an inverted-U dose–response relationship with working memory performance that has no clear parallel in the striatum [DEsposito2015]. Serotonin acting th...”
[DeJong2022] paper:paper-b0ebb9e13d12 “The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different "function" depending on which brain region the reviewer chose to emphasize. Dopamine is a "reward signal" if you read the striatal literature, a "working memory modulator" if you read the prefrontal literature, and a "salience detector" if you read the amygdala literat...”
[Molas2023] paper:paper-b0548de11c5c “The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different "function" depending on which brain region the reviewer chose to emphasize. Dopamine is a "reward signal" if you read the striatal literature, a "working memory modulator" if you read the prefrontal literature, and a "salience detector" if you read the amygdala literat...”
[Morris2025] paper:paper-95c2e1f1250f “The consequence of region-specific reviews — and the literature is dominated by them — is that the same modulatory system appears to have a different "function" depending on which brain region the reviewer chose to emphasize. Dopamine is a "reward signal" if you read the striatal literature, a "working memory modulator" if you read the prefrontal literature, and a "salience detector" if you read the amygdala literat...”
[Marinescu2024] paper:paper-e9e37c821935 “This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei [Marinescu2024, DeSchotten2022, Maciel2026]. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons [Belilos2023, CardozoPinto2025]. The prefrontal cortex r...”
[DeSchotten2022] paper:paper-414dafb1155c “This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei [Marinescu2024, DeSchotten2022, Maciel2026]. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons [Belilos2023, CardozoPinto2025]. The prefrontal cortex r...”
[Maciel2026] paper:paper-fa5d2872a139 “This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei [Marinescu2024, DeSchotten2022, Maciel2026]. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons [Belilos2023, CardozoPinto2025]. The prefrontal cortex r...”
[Belilos2023] paper:paper-89b61251b2cd “This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei [Marinescu2024, DeSchotten2022, Maciel2026]. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons [Belilos2023, CardozoPinto2025]. The prefrontal cortex r...”
[CardozoPinto2025] paper:paper-6985d737857f “This error is compounded when cross-system interactions are neglected. Every cortical and subcortical region receives convergent input from multiple modulatory nuclei [Marinescu2024, DeSchotten2022, Maciel2026]. The nucleus accumbens integrates dopaminergic, serotonergic, cholinergic, and noradrenergic signals on overlapping populations of medium spiny neurons [Belilos2023, CardozoPinto2025]. The prefrontal cortex r...”

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