Activation-synthesis Hypothesis
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The activation-synthesis hypothesis, proposed by
Harvard University Harvard University is a private Ivy League research university in Cambridge, Massachusetts. Founded in 1636 as Harvard College and named for its first benefactor, the Puritan clergyman John Harvard, it is the oldest institution of higher le ...
psychiatrist A psychiatrist is a physician who specializes in psychiatry, the branch of medicine devoted to the diagnosis, prevention, study, and treatment of mental disorders. Psychiatrists are physicians and evaluate patients to determine whether their sy ...
s John Allan Hobson and
Robert McCarley Robert W. McCarley, MD, (1937–2017) was Chair and Professor of Psychiatry at Harvard Medical School and the VA Boston Healthcare System. He is also Director of the Laboratory of Neuroscience located at the Brockton VA Medical Center and the M ...
, is a
neurobiological Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, developme ...
theory of
dream A dream is a succession of images, ideas, emotions, and sensations that usually occur involuntarily in the mind during certain stages of sleep. Humans spend about two hours dreaming per night, and each dream lasts around 5 to 20 minutes, althou ...
s first published in the
American Journal of Psychiatry ''The American Journal of Psychiatry'' is a monthly peer-reviewed medical journal covering all aspects of psychiatry, and is the official journal of the American Psychiatric Association. The first volume was issued in 1844, at which time it was k ...
in December 1977. The differences in neuronal activity of the
brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is cont ...
during waking and
REM sleep Rapid eye movement sleep (REM sleep or REMS) is a unique phase of sleep in mammals and birds, characterized by random rapid movement of the eyes, accompanied by low muscle tone throughout the body, and the propensity of the sleeper to dream viv ...
were observed, and the hypothesis proposes that dreams result from
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
activation during REM sleep. Since then, the hypothesis has undergone an evolution as technology and experimental equipment has become more precise. Currently, a three-dimensional model called AIM Model, described below, is used to determine the different states of the brain over the course of the day and night. The AIM Model introduces a new hypothesis that
primary consciousness Primary consciousness is a term the American biologist Gerald Edelman coined to describe the ability, found in humans and some animals, to integrate observed events with memory to create an awareness of the present and immediate past of the world ar ...
is an important building block on which
secondary consciousness Secondary consciousness is an individual's accessibility to their history and plans. The ability allows its possessors to go beyond the limits of the remembered present of primary consciousness.Edelman, G. M. (2003). Naturalizing consciousness: a ...
is constructed.


Introduction

With the advancement of
brain imaging Neuroimaging is the use of quantitative (computational) techniques to study the neuroanatomy, structure and function of the central nervous system, developed as an objective way of scientifically studying the healthy human brain in a non-invasive ...
technology, the sleep-waking cycle can be studied as never before. The brain can be objectively quantified and identified as being in either one of three states:
awake Wakefulness is a daily recurring brain state and state of consciousness in which an individual is conscious and engages in coherent cognitive and behavioral responses to the external world. Being awake is the opposite of being asleep, in which ...
,
REM sleep Rapid eye movement sleep (REM sleep or REMS) is a unique phase of sleep in mammals and birds, characterized by random rapid movement of the eyes, accompanied by low muscle tone throughout the body, and the propensity of the sleeper to dream viv ...
, and
NREM sleep Non-rapid eye movement sleep (NREM), also known as quiescent sleep, is, collectively, sleep stages 1–3, previously known as stages 1–4. Rapid eye movement sleep (REM) is not included. There are distinct electroencephalographic and other char ...
due to these advanced methods of measurement. It has been shown that global deactivation of the brain from waking state to NREM sleep occurs, and a subsequent reactivation during REM sleep, to a degree greater than during waking.
Consciousness Consciousness, at its simplest, is sentience and awareness of internal and external existence. However, the lack of definitions has led to millennia of analyses, explanations and debates by philosophers, theologians, linguisticians, and scien ...
and its substates,
primary consciousness Primary consciousness is a term the American biologist Gerald Edelman coined to describe the ability, found in humans and some animals, to integrate observed events with memory to create an awareness of the present and immediate past of the world ar ...
and
secondary consciousness Secondary consciousness is an individual's accessibility to their history and plans. The ability allows its possessors to go beyond the limits of the remembered present of primary consciousness.Edelman, G. M. (2003). Naturalizing consciousness: a ...
, play a part in identifying the state of the brain. Primary consciousness is the simple awareness of
perception Perception () is the organization, identification, and interpretation of sensory information in order to represent and understand the presented information or environment. All perception involves signals that go through the nervous system ...
and
emotion Emotions are mental states brought on by neurophysiological changes, variously associated with thoughts, feelings, behavioral responses, and a degree of pleasure or displeasure. There is currently no scientific consensus on a definition. ...
; that is, the awareness of the world via advanced visual and
motor coordination Motor coordination is the orchestrated movement of multiple body parts as required to accomplish intended actions, like walking. This coordination is achieved by adjusting kinematic and kinetic parameters associated with each body part involved in t ...
information your brain receives. Secondary consciousness is an advanced state that includes both primary consciousness and abstract analysis, or thinking, and
metacognitive Metacognition is an awareness of one's thought processes and an understanding of the patterns behind them. The term comes from the root word '' meta'', meaning "beyond", or "on top of".Metcalfe, J., & Shimamura, A. P. (1994). ''Metacognition: knowi ...
components, or the awareness of being aware. Most animals show some stages of primary consciousness, but only humans have been experimentally shown to experience secondary consciousness. The cycle of waking-NREM-REM sleep is essential to mental health of mammals. It has been shown through experimentation that animals subjected to inability to enter REM sleep show an immediate attempt to quickly enter REM stages and long-term effects on motor coordination and habitual motor habits, eventually leading to the death of the animal. It has also been shown that
homeothermic Homeothermy, homothermy or homoiothermy is thermoregulation that maintains a stable internal body temperature regardless of external influence. This internal body temperature is often, though not necessarily, higher than the immediate environmen ...
animals might require sleep to maintain body weight and temperature.


Background


Waking

The waking consciousness is the awareness of the world, our bodies, and ourselves. This includes humans experiencing the awareness of being aware of ourselves, an intrinsic ability to humans. It's the ability to look in a mirror and know that you are looking at yourself, and not just another human being. Being awake allows the distinction between tasks and default brain states, and also distinguishes between background and foreground processing. Being awake allows the person to not only be aware of themselves and the world, but also to have conscious motor coordination and understand the difference between need and want that comes from secondary consciousness.


Difference between sleep and dream

There is a difference between being just asleep and in a state of mind called dreaming. Sleeping can be described as the lack of conscious awareness of the outside world, meaning large portions of the brain that receive and interpret signals are deactivated during this time, while dreaming is a specific state of sleep in which enhanced brain activity has been shown to occur, theorizing the primary consciousness could be active during dreaming. Indeed, during dreams we are consciously aware of our surroundings, and assuredly have a certain perception and emotion throughout the course of the dream, suggesting that at least part of the primary consciousness is activated during the dream.


Dream

A dream has all features of primary consciousness but is produced in the brain without external stimulation. Unlike the waking state, the brain cannot recognize its own condition; that it is in the midst of the dream and is not the same as the real world. The brain has a single-minded state of primary consciousness during dreaming, which allows the brain to reach greater perception and awareness of a single scenario out of images and dreams. This is called the dream consciousness.


Four stages of sleep

The four
sleep Sleep is a sedentary state of mind and body. It is characterized by altered consciousness, relatively inhibited sensory activity, reduced muscle activity and reduced interactions with surroundings. It is distinguished from wakefulness by a de ...
stages have been identified as follows: sleep onset stage I, late-night stage II, and deep sleep stages III and IV. Deep sleep stages III and IV all occur during the first half of the night, while lighter stages I and II occur during the later half. During standard sleep laboratory measurements, the states of sleep and waking have
behavioral Behavior (American English) or behaviour (British English) is the range of actions and mannerisms made by individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as well ...
, polygraphic, and
psychological Psychology is the scientific study of mind and behavior. Psychology includes the study of conscious and unconscious phenomena, including feelings and thoughts. It is an academic discipline of immense scope, crossing the boundaries between t ...
manifestation within the pontine brainstem. These states are regulated by a reciprocal relationship between two types of neuronal cells, aminergic inhibitory cells such as
serotonin Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and vas ...
and
norepinephrine Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad'', ...
and
cholinergic Cholinergic agents are compounds which mimic the action of acetylcholine and/or butyrylcholine. In general, the word "choline" describes the various quaternary ammonium salts containing the ''N'',''N'',''N''-trimethylethanolammonium cation. F ...
excitatory cells such as
acetylcholine Acetylcholine (ACh) is an organic chemical that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Part ...
. Changes in the sleep stages occur when the activity curves of these neurons cross. REM sleep stage I is a state of sleep just above and most closely linked to sleep onset stage I.


NREM

NREM sleep can be described as the stages of sleep that show greatly decreased brain activity. There are four different stages of NREM sleep. The brain shows dulled or limited senses of perception, though the thought process has been shown to be logical and perseverative. Episodic movements of the body occur during these stages, though they are involuntary movements.


REM

REM sleep is an evolutionarily recent behavior of humans. REM stands for rapid eye movement. It is the deepest sleep a mammal can go into. It is regulated by the pontine brainstem. Infants spend most of their time in REM sleep, and rather than enter stage 1 sleep they go directly to REM sleep. Most REM sleep occurs just above stage I of sleep, and experiences different mental abilities than during NREM sleep. The thought process is non-logical and often bizarre, sensation and perception is vivid but created internally by the brain, and the body's movements are inhibited. Most REM stages last 10–15 minutes, and the average human will go through 4–6 of these stages during sleep each night. Subsequent REM stages increase in duration, so the last REM stage before awakening is the longest and most vivid. During REM sleep the brain shows increased states of minimal inhibition, which degrades in our ability to recognize the state for which it is; a dream. It has been proposed that REM sleep is necessary for preparation of many integrative functions, of which one is consciousness. It supports the idea that sleep, and dreaming, is necessary preparation for the next day's processes. The scientific tracking of REM sleep stages can be measured by neuronal signals within the pontine brainstem. The interactions of aminergic inhibitory neurons and cholinergic excitatory neurons can be measured, and REM sleep occurs when aminergic cells are at their least active and cholinergic cells are at their most active.


Evolution of REM

It has been stated that REM sleep is a recent evolutionary behavior in homeothermic animals. In both, there is increased REM sleep in the early stages of life. In humans, REM sleep peaks during the third trimester of gestation, and quickly falls after birth as primary consciousness declines and secondary consciousness grows with the development of the brain. The developing control over stages of sleep and waking suggests that sleep and REM has developed as a way to self-activate in order to anticipate awake-state circumstances.


Neuronic modeling

Within the
pons The pons (from Latin , "bridge") is part of the brainstem that in humans and other bipeds lies inferior to the midbrain, superior to the medulla oblongata and anterior to the cerebellum. The pons is also called the pons Varolii ("bridge of Va ...
, the modeling and tracking of these aminergic inhibitory neurons and cholinergic excitatory neurons occurs via the study of
PGO waves Ponto-geniculo-occipital waves or PGO waves are distinctive wave forms of propagating activity between three key brain regions: the pons, lateral geniculate nucleus, and occipital lobe; specifically, they are phasic field potentials. These waves c ...
. These are phasic waves that occur in cycles, and originate from the pontine brainstem (P),
lateral geniculate In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projec ...
of
thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
(G), and
occipital cortex The occipital lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The name derives from its position at the back of the head, from the Latin ''ob'', "behind", and ''caput'', "head". The occipital lobe is the vi ...
(O). Aminergic
monoamines Monoamine neurotransmitters are neurotransmitters and neuromodulators that contain one amino group connected to an aromatic ring by a two-carbon chain (such as -CH2-CH2-). Examples are dopamine, norepinephrine and serotonin. All monoamines ar ...
serotonin,
noradrenaline Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad'', ...
,
histamine Histamine is an organic nitrogenous compound involved in local immune responses, as well as regulating physiological functions in the gut and acting as a neurotransmitter for the brain, spinal cord, and uterus. Since histamine was discovered in ...
, and
dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic compound, organic chemical of the catecholamine and phenethylamine families. Dopamine const ...
are balanced between acetylcholine cholinergic signals, and play a part in the regulation of cognition. Aminergic cell signal strength is lowest during REM sleep, increases during NREM, and is highest at waking. Cholinergic cell signal strength is highest during REM, declines during NREM, and is lowest at waking. Changes in sleep state and phase occur when two activity paths cross.


Theory

The development of consciousness is a gradual, time-consuming and lifelong process that builds upon and uses a more primitive virtual reality generator that is more definable in our dreams. As such, the development of secondary consciousness during the lifetime requires a blank consciousness that during REM sleep creates an imaginary self that has movements and experiences emotions. This is an experimental state not associated with awareness, and this state, or protoconscious, is able to be reached during childhood. This protoconsciousness is a protoself created early in life by the brain as a building block for consciousness to develop, and provides intrinsic predictions of external inputs created by dreaming.


Original activation-synthesis hypothesis model

Hobson and McCarley originally proposed in the 1970s that the differences in the waking-NREM-REM sleep cycle was the result of interactions between aminergic REM-off cells and cholinergic REM-on cells. This was perceived as the activation-synthesis model, stating that brain activation during REM sleep results in synthesis of dream creation. Hobson's five cardinal characteristics include: intense emotions, illogical content, apparent sensory impressions, uncritical acceptance of dream events, and difficulty in being remembered.


Current model – AIM

Thanks to the development of technology since the original proposal, new experimental data has been collected and additional mechanistic details of neuronal control have been developed. It has been determined that consciousness states can be described with three values, and the AIM model is a model that uses these values for representing the similarities and differences between waking and dreaming. It is a three-dimensional
state-space A state space is the set of all possible configurations of a system. It is a useful abstraction for reasoning about the behavior of a given system and is widely used in the fields of artificial intelligence and game theory. For instance, the toy ...
model that describes different states of the brain and their variance throughout the day and night. It is composed of three different values: A – activation, I – input-output gating, and M – modulation. The model is limited however, in that it cannot yet explain the regional differences in brain activity that distinguish REM sleep from waking. Other limitations include the inability to quantifiably identify and measure M in humans. During waking and activation of primary and secondary consciousnesses, high values of A, I, and M have been observed, but during REM sleep high values of A but low I and M have been observed.


Protoconsciousness

The protoconsciousness is template of consciousness that occurs during sleep, and on which can be constructed other mental conscious processes. Early in childhood, it has been said that this protoconsciousness is where secondary aspects of consciousness are originally developed and tested by the primary consciousness, and the person can slowly develop increased secondary consciousness throughout their life as their protoconscious template is further expanded, developed, and creates more vivid ideas and representations of secondary consciousness.


Activation (A)

Large parts of the brain that are activated and sending signals during waking are inactive during NREM sleep and become reactivated during REM sleep. It is based on the fact that the brain and its neural circuitry is plastic and self-regulating, especially in its own activation and inactivation. This was observed by two experiments: development of sleepiness after dopamine neuron destruction in
substantia nigra The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra app ...
in the
midbrain The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', " ...
, and discovery of the
reticular activating system The reticular formation is a set of interconnected nuclei that are located throughout the brainstem. It is not anatomically well defined, because it includes neurons located in different parts of the brain. The neurons of the reticular formatio ...
, which are visual cues received through our eyes and to our brain that begin the waking process, that waking consciousness depends sleep. Following these studies, it became clear that activity levels and quality of consciousness were functions of brain activation and deactivation.


Input-output gating (I)

It has been shown that the internal activation of the brain is associated with the inhibition of both external sensory input and motor output. This implies that the brain is actively kept offline during REM, and the brainstem guarantees the coordination of factors I and A via the input-output gate control within the brainstem. PGO waves play a part in the ability of the brain to remain asleep while constituting the building blocks for perception and fine motor control via their phasic coordination. It has therefore been proposed that PGO signals are used in the construction of visual imagery of dreams.


Modulation (M)

The neuromodulator release of aminergic neurons have a broad chemical influence on the brain; they instruct other neurons to keep or discard a record of information they've processed. The mechanics of modulation are not known at this time, and modulation has yet to be quantitatively identified. Qualitatively, aminergic modulation has been shown to be strong during waking but lower during sleep, but more studies need to be conducted. Numerous studies have emerged from the discipline of
computational neuroscience Computational neuroscience (also known as theoretical neuroscience or mathematical neuroscience) is a branch of neuroscience which employs mathematical models, computer simulations, theoretical analysis and abstractions of the brain to u ...
that support to the AIM model. The theory of
Metalearning Meta-learning is a branch of metacognition concerned with learning about one's own learning and learning processes. The term comes from the meta prefix's modern meaning of an abstract recursion, or "X about X", similar to its use in metaknowl ...
in particular describes how these neuromodulators facilitate dynamic learning, though a series of interpretive models all consistent with the AIM model.


Implications

The three-dimensional AIM model shows that during the cycle of brain states waking-NREM-REM, the brain is dynamically changing constantly, and that this state space described by the AIM has an infinite number of subregions other than the main three. It proposes that via a protoconsciousness brain activation during sleep is necessary for the development and maintenance of waking consciousness and other higher-order brain functions such as problem solving. It suggests the possibility that the state of waking consciousness is only present in humans due to the evolution of extensive cortical structures within the brain. Dreaming is a state of the brain that is similar to yet different from the waking consciousness, and interaction and correlation between the two is necessary for optimal performance from both. One study conducted measuring brain activity via
EEG Electroencephalography (EEG) is a method to record an electrogram of the spontaneous electrical activity of the brain. The biosignals detected by EEG have been shown to represent the postsynaptic potentials of pyramidal neurons in the neocortex ...
used Hobson's AIM model to show that quantitatively dream consciousness is remarkably similar to waking consciousness.


References

{{Dreaming Sleep physiology Dream Unsolved problems in neuroscience Consciousness