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Friday, June 15, 2018

Myelin

From Wikipedia, the free encyclopedia

Structure of a typical neuron
Myelin sheath

Myelin is a lipid-rich substance that surrounds the axon of some nerve cells, forming an electrically insulating layer. It is essential for the proper functioning of the nervous system. It is produced by specialized glial cells through extension of their cellular processes.

The production of the myelin sheath is called myelination or myelinogenesis. In humans, myelination begins early in the 3rd trimester,[1] although little myelin exists in the brain at the time of birth. During infancy, myelination occurs quickly, leading to a child's fast development, including crawling and walking in the first year. Myelination continues through the adolescent stage of life.

Schwann cells supply the myelin for the peripheral nervous system, whereas oligodendrocytes, specifically of the interfascicular type, myelinate the axons of the central nervous system. Myelin is considered a defining characteristic of the (gnathostome) vertebrates, but myelin-like sheaths have also been seen in some invertebrates,[2][3] although they are quite different from vertebrate myelin at molecular level. Myelin was discovered in 1854 by Rudolf Virchow.[4]

Composition


Transmission electron micrograph of a cross-section of a myelinated axon, generated at the Electron Microscopy Facility at Trinity College, Hartford CT

Myelin comprises different cell types and varies in chemical composition and configuration but performs the same insulating function. Myelinated axons are white; hence, the "white matter" of the brain. Myelin insulates axons from electrically charged atoms and molecules. These charged particles (ions) are found in the fluid surrounding the entire nervous system. Under a microscope, myelin looks like strings of sausages.

Cholesterol is an essential component of myelin,[5] which comprises about 40% water; the dry mass comprises between 60% and 75% lipids and between 15% and 25% proteins. Myelin basic protein (MBP) constitutes ~23% of myelin protein,[6] myelin oligodendrocyte glycoprotein, and proteolipid protein (PLP, which makes up ~50% of myelin protein[7]). The primary lipid of myelin is a glycolipid called galactocerebroside. The intertwining hydrocarbon chains of sphingomyelin strengthen the myelin sheath. In brain, the myelin sheath covers the fibers of the corpus callosum, which constitute the inner part of the cerebral hemisphere.

Function


Action potential propagation in myelinated neurons is faster than in unmyelinated neurons because of Saltatory conduction.

Cross section of a myelinated axon
1. Axon
2. Nucleus of Schwann Cell
3. Schwann Cell
4. Myelin Sheath
5. Neurilemma

The main purpose of a myelin sheath is to increase the speed at which impulses propagate along the myelinated fiber. Along unmyelinated fibers, impulses continuously move as waves, but, in myelinated fibers, they "hop" or propagate by saltatory conduction. Myelin decreases capacitance and increases electrical resistance across the cell membrane (the axolemma). Thus, myelination prevents the electric current from leaving the axon. It has been suggested that myelin permits larger body size by maintaining agile communication between distant body parts.[2]

Myelinated fibers lack voltage-gated ion channels (approximately 25 μm−2) along the myelinated internodes, exposing them only at the nodes of Ranvier. Here, they are found far more abundantly (between 2,000 and 12,000 μm−2).[8] Myelinated fibers succeed in reducing sodium leakage into the extracellular fluid (ECF), maintaining a strong separation of charge between the intracellular fluid (ICF) and the ECF. This increases sodium's ability to travel along the axon more freely. However, the sodium diffuses along the axolemma rapidly, but is decremental by nature. The sodium cannot trigger the opening of the voltage-gated sodium channels as it becomes weaker. The nodes of Ranvier, being exposed to the ECF every 1 mm or so, contain large amounts of voltage-gated sodium channels, and allow enough sodium into the axon to regenerate the action potential.[9] Each time the action potential reaches a node of Ranvier, it is restored to its original action potential (+35 mV).[8]

When a peripheral fiber is severed, the myelin sheath provides a track along which regrowth can occur. However, the myelin layer does not ensure a perfect regeneration of the nerve fiber. Some regenerated nerve fibers do not find the correct muscle fibers, and some damaged motor neurons of the peripheral nervous system die without regrowth. Damage to the myelin sheath and nerve fiber is often associated with increased functional insufficiency.

Unmyelinated fibers and myelinated axons of the mammalian central nervous system do not regenerate.

Some studies have revealed that optic nerve fibers can be regenerated in postnatal rats. This regeneration depends upon two conditions: axonal die-back has to be prevented with appropriate neurotrophic factors, and neurite growth inhibitory components have to be inactivated. These studies may lead to further understanding of nerve fiber regeneration in the central nervous system.[citation needed]

Disorders

Demyelination

Demyelination is the loss of the myelin sheath insulating the nerves, and is the hallmark of some neurodegenerative autoimmune diseases, including multiple sclerosis, acute disseminated encephalomyelitis, neuromyelitis optica, transverse myelitis, chronic inflammatory demyelinating polyneuropathy, Guillain–Barré syndrome, central pontine myelinosis, inherited demyelinating diseases such as leukodystrophy, and Charcot-Marie-Tooth disease. Sufferers of pernicious anaemia can also suffer nerve damage if the condition is not diagnosed quickly. Subacute combined degeneration of spinal cord secondary to pernicious anaemia can lead to slight peripheral nerve damage to severe damage to the central nervous system, affecting speech, balance, and cognitive awareness. When myelin degrades, conduction of signals along the nerve can be impaired or lost, and the nerve eventually withers.[clarification needed] A more serious case of myelin deterioration is called Canavan disease.

The immune system may play a role in demyelination associated with such diseases, including inflammation causing demyelination by overproduction of cytokines via upregulation of tumor necrosis factor[10] or interferon.

Symptoms

Demyelination results in diverse symptoms determined by the functions of the affected neurons. It disrupts signals between the brain and other parts of the body; symptoms differ from patient to patient, and have different presentations upon clinical observation and in laboratory studies.

Typical symptoms include:
  • blurriness in the central visual field that affects only one eye, may be accompanied by pain upon eye movement
  • double vision
  • loss of vision/hearing
  • odd sensation in legs, arms, chest, or face, such as tingling or numbness (neuropathy)
  • weakness of arms or legs
  • cognitive disruption, including speech impairment and memory loss
  • heat sensitivity (symptoms worsen or reappear upon exposure to heat, such as a hot shower)
  • loss of dexterity
  • difficulty coordinating movement or balance disorder
  • difficulty controlling bowel movements or urination
  • fatigue
  • tinnitus[11]

Myelin repair

Research to repair damaged myelin sheaths is ongoing. Techniques include surgically implanting oligodendrocyte precursor cells in the central nervous system and inducing myelin repair with certain antibodies. While results in mice have been encouraging (via stem cell transplantation), whether this technique can be effective in replacing myelin loss in humans is still unknown.[12] Cholinergic treatments, such as acetylcholinesterase inhibitors (AChEIs), may have beneficial effects on myelination, myelin repair, and myelin integrity. Increasing cholinergic stimulation also may act through subtle trophic effects on brain developmental processes and particularly on oligodendrocytes and the lifelong myelination process they support. By increasing oligodendrocyte cholinergic stimulation, AChEIs, and other cholinergic treatments, such as nicotine, possibly could promote myelination during development and myelin repair in older age.[13] Glycogen synthase kinase 3β inhibitors such as lithium chloride have been found to promote myelination in mice with damaged facial nerves.[14] Cholesterol is a necessary nutrient for the myelin sheath.

Dysmyelination

Dysmyelination is characterized by a defective structure and function of myelin sheaths; unlike demyelination, it does not produce lesions. Such defective sheaths often arise from genetic mutations affecting the biosynthesis and formation of myelin. The shiverer mouse represents one animal model of dysmyelination. Human diseases where dysmyelination has been implicated include leukodystrophies (Pelizaeus–Merzbacher disease, Canavan disease, phenylketonuria) and schizophrenia.[15][16][17]

Invertebrate myelin

Functionally equivalent myelin-like sheaths are found in several invertebrate taxa including Oligochaete, Penaeid, Palaemonid, and Calanoids. These myelin-like sheaths share several structural features with the sheaths found in vertebrates including multiplicity of membranes, condensation of membrane, and nodes.[2] However, the nodes in vertebrates are annular; i.e. they encircle the axon. In contrast, nodes found in the sheaths of invertebrates are either annular or fenestrated; i.e. they are restricted to "spots." It is notable that the fastest recorded conduction speed (across both vertebrates and invertebrates) is found in the ensheathed axons of the Kuruma shrimp, an invertebrate,[2] ranging between 90 and 200 m/s[3] (cf 100-120 m/s for the fastest myelinated vertebrate axon.)

Neuroglia

From Wikipedia, the free encyclopedia

Neuroglia
Glial Cell Types.png
Illustration of the four different types of glial cells found in the central nervous system: ependymal cells (light pink), astrocytes (green), microglial cells (dark red), and oligodendrocytes (light blue).
Details
Precursor Neuroectoderm for macroglia, and hematopoietic stem cells for microglia
System Nervous system
Identifiers
MeSH D009457
TA A14.0.00.005
TH H2.00.06.2.00001
FMA 54541
Anatomical terms of microanatomy

Neuroglia, also called glial cells or simply glia, are non-neuronal cells in the central nervous system (brain and spinal cord) and the peripheral nervous system. They maintain homeostasis, form myelin, and provide support and protection for neurons.[1] In the central nervous system, glial cells include oligodendrocytes, astrocytes, ependymal cells, and microglia, and in the peripheral nervous system glial cells include Schwann cells and satellite cells. They have four main functions: (1) To surround neurons and hold them in place (2) To supply nutrients and oxygen to neurons (3) To insulate one neuron from another (4) To destroy pathogens and remove dead neurons. They also play a role in neurotransmission and synaptic connections,[2] and in physiological processes like breathing.[3][4] .

Glia were discovered in 1856, by the pathologist Rudolf Virchow in his search for a "connective tissue" in the brain.[5] The term derives from Greek γλία and γλοία "glue"(/ˈɡlə/ or /ˈɡlə/), and suggests the original impression that they were the glue of the nervous system.

Types


Neuroglia of the brain shown by Golgi's method

Astrocytes can be identified in culture because, unlike other mature glia, they express glial fibrillary acidic protein (GFAP)

Glial cells in a rat brain stained with an antibody against GFAP

Different types of neuroglia

Macroglia

Derived from ectodermal tissue.

Location Name Description
CNS Astrocytes The most abundant type of macroglial cell in the CNS,[6] astrocytes (also called astroglia) have numerous projections that link neurons to their blood supply while forming the blood-brain barrier. They regulate the external chemical environment of neurons by removing excess potassium ions, and recycling neurotransmitters released during synaptic transmission. Astrocytes may regulate vasoconstriction and vasodilation by producing substances such as arachidonic acid, whose metabolites are vasoactive.

Astrocytes signal each other using ATP. The gap junctions (also known as electrical synapses) between astrocytes allow the messenger molecule IP3 to diffuse from one astrocyte to another. IP3 activates calcium channels on cellular organelles, releasing calcium into the cytoplasm. This calcium may stimulate the production of more IP3 and cause release of ATP through channels in the membrane made of pannexins. The net effect is a calcium wave that propagates from cell to cell. Extracellular release of ATP, and consequent activation of purinergic receptors on other astrocytes, may also mediate calcium waves in some cases.

In general, there are two types of astrocytes, protoplasmic and fibrous, similar in function but distinct in morphology and distribution. Protoplasmic astrocytes have short, thick, highly branched processes and are typically found in gray matter. Fibrous astrocytes have long, thin, less branched processes and are more commonly found in white matter.

It has recently been shown that astrocyte activity is linked to blood flow in the brain, and that this is what is actually being measured in fMRI.[7] They also have been involved in neuronal circuits playing an inhibitory role after sensing changes in extracellular calcium.[8]
CNS Oligodendrocytes Oligodendrocytes are cells that coat axons in the central nervous system (CNS) with their cell membrane, forming a specialized membrane differentiation called myelin, producing the myelin sheath. The myelin sheath provides insulation to the axon that allows electrical signals to propagate more efficiently.[9]
CNS Ependymal cells Ependymal cells, also named ependymocytes, line the spinal cord and the ventricular system of the brain. These cells are involved in the creation and secretion of cerebrospinal fluid (CSF) and beat their cilia to help circulate the CSF and make up the blood-CSF barrier. They are also thought to act as neural stem cells.[10]
CNS Radial glia Radial glia cells arise from neuroepithelial cells after the onset of neurogenesis. Their differentiation abilities are more restricted than those of neuroepithelial cells. In the developing nervous system, radial glia function both as neuronal progenitors and as a scaffold upon which newborn neurons migrate. In the mature brain, the cerebellum and retina retain characteristic radial glial cells. In the cerebellum, these are Bergmann glia, which regulate synaptic plasticity. In the retina, the radial Müller cell is the glial cell that spans the thickness of the retina and, in addition to astroglial cells,[11] participates in a bidirectional communication with neurons.[12]
PNS Schwann cells Similar in function to oligodendrocytes, Schwann cells provide myelination to axons in the peripheral nervous system (PNS). They also have phagocytotic activity and clear cellular debris that allows for regrowth of PNS neurons.[13]
PNS Satellite cells Satellite glial cells are small cells that surround neurons in sensory, sympathetic, and parasympathetic ganglia.[14] These cells help regulate the external chemical environment. Like astrocytes, they are interconnected by gap junctions and respond to ATP by elevating intracellular concentration of calcium ions. They are highly sensitive to injury and inflammation, and appear to contribute to pathological states, such as chronic pain.[15]
PNS Enteric glial cells Are found in the intrinsic ganglia of the digestive system. They are thought to have many roles in the enteric system, some related to homeostasis and muscular digestive processes.[16]

Microglia

Microglia are specialized macrophages capable of phagocytosis that protect neurons of the central nervous system.[17] They are derived from the earliest wave of mononuclear cells that originate in yolk sac blood islands early in development, and colonize the brain shortly after the neural precursors begin to differentiate.[18]

These cells are found in all regions of the brain and spinal cord. Microglial cells are small relative to macroglial cells, with changing shapes and oblong nuclei. They are mobile within the brain and multiply when the brain is damaged. In the healthy central nervous system, microglia processes constantly sample all aspects of their environment (neurons, macroglia and blood vessels). In a healthy brain, microglia direct the immune response to brain damage and play an important role in the inflammation that accompanies the damage. Many diseases and disorders are associated with deficient microglia, such as Alzheimer's disease, Parkinson's disease, and ALS.

Other

Pituicytes from the posterior pituitary are glia cells with characteristics in common to astrocytes.[19] Tanycytes in the median eminence of the hypothalamus are a type of ependymal cell that descend from radial glia and line the base of the third ventricle.[20]

Total number

In general, neuroglial cells are smaller than neurons; there are about 86 billion neurons and 85 billion "non-neuronal" (glia) cells in the human male brain. Glial cells make up about half the total volume of the brain and spinal cord.[21][22]^ The ratio varies from one part of the brain to another. The glia/neuron ratio in the cerebral cortex is 3.72 (60.84 billion glia (72%); 16.34 billion neurons), while that of the cerebellum is only 0.23 (16.04 billion glia; 69.03 billion neurons). The ratio in the cerebral cortex gray matter is 1.48, with 3.76 for the gray and white matter combined.[21] The ratio of the basal ganglia, diencephalon and brainstem combined is 11.35.[21]

Most cerebral cortex glia are oligodendrocytes (75.6%); astrocytes account for 17.3% and microglia for 6.5%.[23]

Development


23-week fetal brain culture astrocyte

Most glia are derived from ectodermal tissue of the developing embryo, in particular the neural tube and crest. The exception is microglia, which are derived from hemopoietic stem cells. In the adult, microglia are largely a self-renewing population and are distinct from macrophages and monocytes, which infiltrate the injured and diseased CNS.

In the central nervous system, glia develop from the ventricular zone of the neural tube. These glia include the oligodendrocytes, ependymal cells, and astrocytes. In the peripheral nervous system, glia derive from the neural crest. These PNS glia include Schwann cells in nerves and satellite glial cells in ganglia.

Current research involving glial cells in the human cochlea proposes that these cells are the common precursor to both mature Schwann cells and satellite glial cells. Additionally, the peripheral glial cells located along the peripheral processes expressed NGFR, indicating a phenotype distinct from the peripheral glial cells located along the central processes.[24]

Capacity to divide

Glia retain the ability to undergo cell division in adulthood, whereas most neurons cannot. The view is based on the general deficiency of the mature nervous system in replacing neurons after an injury, such as a stroke or trauma, while very often there is a profound proliferation of glia, or gliosis near or at the site of damage. However, detailed studies found no evidence that 'mature' glia, such as astrocytes or oligodendrocytes, retain the ability of mitosis. Only the resident oligodendrocyte precursor cells seem to keep this ability after the nervous system matures. On the other hand, there are a few regions in the mature nervous system, such as the dentate gyrus of the hippocampus and the subventricular zone, where generation of new neurons can be observed.[25]

Glial cells are known to be capable of mitosis. By contrast, scientific understanding of whether neurons are permanently post-mitotic,[26] or capable of mitosis,[27][28][29] is still developing. In the past, glia had been considered[by whom?] to lack certain features of neurons. For example, glial cells were not believed to have chemical synapses or to release transmitters. They were considered to be the passive bystanders of neural transmission. However, recent studies have shown this to be untrue.[30]

Functions

Some glial cells function primarily as the physical support for neurons. Others regulate the internal environment of the brain, especially the fluid surrounding neurons and their synapses, and nutrify neurons. During early embryogenesis, glial cells direct the migration of neurons and produce molecules that modify the growth of axons and dendrites.

Neuron repair and development

Glia are also crucial in the development of the nervous system and in processes such as synaptic plasticity and synaptogenesis. Glia have a role in the regulation of repair of neurons after injury. In the central nervous system (CNS), glia suppress repair. Glial cells known as astrocytes enlarge and proliferate to form a scar and produce inhibitory molecules that inhibit regrowth of a damaged or severed axon. In the peripheral nervous system (PNS), glial cells known as Schwann cells promote repair. After axonal injury, Schwann cells regress to an earlier developmental state to encourage regrowth of the axon. This difference between the CNS and the PNS, raises hopes for the regeneration of nervous tissue in the CNS. For example, a spinal cord may be able to be repaired following injury or severance. Schwann cells are also known as neuri-lemmocytes. These cells envelop nerve fibers of the PNS by winding repeatedly around a nerve fiber with the nucleus inside of it. This process creates a myelin sheath, which not only aids in conductivity but also assists in the regeneration of damaged fibers.

Myelin sheath creation

Oligodendrocytes are another type of glial cell of the CNS. These dendrocytes resemble an octopus bulbous body and contain up to fifteen arm-like processes. Each “arm” reaches out to a nerve fiber and spirals around it, creating a myelin sheath. This myelin sheath insulates the nerve fiber from the extracellular fluid as well as speeds up the signal conduction in the nerve fiber.[31]

Neurotransmission

Recent research indicates that glial cells of the hippocampus and cerebellum participate in synaptic transmission, regulate the clearance of neurotransmitters from the synaptic cleft, and release gliotransmitters such as ATP, which modulate synaptic function.[32]

Astrocytes are crucial in clearance of neurotransmitters from within the synaptic cleft, which provides distinction between arrival of action potentials and prevents toxic build-up of certain neurotransmitters such as glutamate (excitotoxicity). It is also thought that glia play a role in many neurological diseases, including Alzheimer's disease.[33] Furthermore, at least in vitro, astrocytes can release gliotransmitter glutamate in response to certain stimulation. Another unique type of glial cell, the oligodendrocyte precursor cells or OPCs, have very well-defined and functional synapses from at least two major groups of neurons.[34] The only notable differences between neurons and glial cells are neurons' possession of axons and dendrites, and capacity to generate action potentials.

Clinical significance


Neoplastic glial cells stained with an antibody against GFAP (brown), from a brain biopsy

While glial cells in the PNS frequently assist in regeneration of lost neural functioning, loss of neurons in the CNS does not result in a similar reaction from neuroglia.[13] In the CNS, regrowth will only happen if the trauma was mild, and not severe.[35] When severe trauma presents itself, the survival of the remaining neurons becomes the optimal solution. However, some studies investigating the role of glial cells in Alzheimer's Disease are beginning to contradict the usefulness of this feature, and even claim it can "exacerbate" the disease.[36] In addition to impacting the potential repair of neurons in Alzheimer's Disease, scarring and inflammation from glial cells have been further implicated in the degeneration of neurons caused by Amyotrophic lateral sclerosis.[37]

In addition to neurodegenerative diseases, a wide range of harmful exposure, such as hypoxia, or physical trauma, can lead to the end result of physical damage to the CNS.[35] Generally, when damage occurs to the CNS, glial cells cause Apoptosis among the surrounding cellular bodies.[35] Then, there is a large amount of microglial activity, which results in inflammation, and finally, there is a heavy release of growth inhibiting molecules.[35]

History

Glia were first described in 1856 by the pathologist Rudolf Virchow in a comment to his 1846 publication on connective tissue. A more detailed description of glial cells was provided in the 1858 book Cellular Pathology by the same author.[38]

When markers for different types of cells were analyzed, Einstein's brain was discovered to contain significantly more glia than normal brains in the left angular gyrus, an area thought to be responsible for mathematical processing and language.[39]

The ratio of glia to neurons increases with our definition of intelligence. Not only does the ratio of glia to neurons increase through evolution, but so does the size of the glia. Astroglial cells in the human have a volume 27 times greater than the same cells in the mouse's brain.[40]

These important scientific findings may begin to shift the neuron-specific perspective into a more holistic view of the brain which encompasses the glial cells as well. The glia's importance is becoming ever more clear as time goes on and new research is conducted. For most of the last century, scientists had written off glial cells as being nothing more than the structure and foundations that hold the neurons in place. But now, there is direct evidence that correlates the number of glial cells in the brain with the amount of intelligence that any given species possesses.[41] Future research will begin to shed light on the mysterious, yet increasingly crucial, role of glial cells.

Reductionism

From Wikipedia, the free encyclopedia

Descartes claimed that non-human animals could be explained reductively as automata — De homine, 1662.

Reductionism is any of several related philosophical ideas regarding the associations between phenomena which can be described in terms of other simpler or more fundamental phenomena.[1]

The Oxford Companion to Philosophy suggests that reductionism is "one of the most used and abused terms in the philosophical lexicon" and suggests a three part division:[2]
  1. Ontological reductionism: a belief that the whole of reality consists of a minimal number of parts.
  2. Methodological reductionism: the scientific attempt to provide explanation in terms of ever smaller entities.
  3. Theory reductionism: the suggestion that a newer theory does not replace or absorb an older one, but reduces it to more basic terms. Theory reduction itself is divisible into three parts: translation, derivation and explanation.[3]
Reductionism can be applied to any phenomenon, including objects, explanations, theories, and meanings.[3][4] [5]

For the sciences, application of methodological reductionism attempts explanation of entire systems in terms of their individual, constituent parts and their interactions. For example, the temperature of a gas is reduced to nothing but the average kinetic energy of its molecules in motion. Thomas Nagel speaks of psychophysical reductionism (the attempted reduction of psychological phenomena to physics and chemistry), as do others and physico-chemical reductionism (the attempted reduction of biology to physics and chemistry), again as do others.[6] In a very simplified and sometimes contested form, such reductionism is said to imply that a system is nothing but the sum of its parts.[4][7] However, a more nuanced opinion is that a system is composed entirely of its parts, but the system will have features that none of the parts have.[8] "The point of mechanistic explanations is usually showing how the higher level features arise from the parts."[7]

Other definitions are used by other authors. For example, what John Polkinghorne terms conceptual or epistemological reductionism[4] is the definition provided by Simon Blackburn[9] and by Jaegwon Kim:[10] that form of reductionism concerning a program of replacing the facts or entities entering statements claimed to be true in one type of discourse with other facts or entities from another type, thereby providing a relationship between them. Such an association is provided where the same idea can be expressed by "levels" of explanation, with higher levels reducible if need be to lower levels. This use of levels of understanding in part expresses our human limitations in remembering detail. However, "most philosophers would insist that our role in conceptualizing reality [our need for an hierarchy of "levels" of understanding] does not change the fact that different levels of organization in reality do have different properties."[8]

Reductionism strongly represents a certain perspective of causality. In a reductionist framework, the phenomena that can be explained completely in terms of relations between other more fundamental phenomena, are termed epiphenomena. Often there is an implication that the epiphenomenon exerts no causal agency on the fundamental phenomena that explain it. The epiphenomena are sometimes said to be "nothing but" the outcome of the workings of the fundamental phenomena, although the epiphenomena might be more clearly and efficiently described in very different terms. There is a tendency to avoid considering an epiphenomenon as being important in its own right. This attitude may extend to cases where the fundamentals are not obviously able to explain the epiphenomena, but are expected to by the speaker. In this way, for example, morality can be deemed to be "nothing but" evolutionary adaptation, and consciousness can be considered "nothing but" the outcome of neurobiological processes.

Reductionism should be distinguished from eliminationism: reductionists do not deny the existence of phenomena, but explain them in terms of another reality; eliminationists deny the existence of the phenomena themselves. For example, eliminationists deny the existence of life by their explanation in terms of physical and chemical processes.

Reductionism also does not preclude the existence of what might be termed emergent phenomena, but it does imply the ability to understand those phenomena completely in terms of the processes from which they are composed. This reductionist understanding is very different from emergentism, which intends that what emerges in "emergence" is more than the sum of the processes from which it emerges.[11]

Types

Most philosophers delineate three types of reductionism and anti-reductionism.[2]

Ontological reductionism

Ontological reductionism is the belief that reality is composed of a minimum number of kinds of entities or substances. This claim is usually metaphysical, and is most commonly a form of monism, in effect claiming that all objects, properties and events are reducible to a single substance. (A dualist who is an ontological reductionist would believe that everything is reducible to two substances — as one possible example, a dualist might claim that reality is composed of "matter" and "spirit".)

Richard Jones divides ontological reductionism into two: the reductionism of substances (e.g., the reduction of mind to matter) and the reduction of the number of structures operating in nature (e.g., the reduction of one physical force to another). This permits scientists and philosophers to affirm the former while being anti-reductionists regarding the latter.[12]

Nancey Murphy has claimed that there are two species of ontological reductionism: one that denies that wholes are anything more than their parts; and the stronger thesis of atomist reductionism that wholes are not "really real". She admits that the phrase "really real" is apparently senseless but nonetheless has tried to explicate the supposed difference between the two.[13]

Ontological reductionism denies the idea of ontological emergence, and claims that emergence is an epistemological phenomenon that only exists through analysis or description of a system, and does not exist fundamentally.[14]

Ontological reductionism takes two different forms: token ontological reductionism and type ontological reductionism.

Token ontological reductionism is the idea that every item that exists is a sum item. For perceivable items, it affirms that every perceivable item is a sum of items with a lesser degree of complexity. Token ontological reduction of biological things to chemical things is generally accepted.

Type ontological reductionism is the idea that every type of item is a sum type of item, and that every perceivable type of item is a sum of types of items with a lesser degree of complexity. Type ontological reduction of biological things to chemical things is often rejected.[15]

Michael Ruse has criticized ontological reductionism as an improper argument against vitalism.[16]

Methodological reductionism

Methodological reductionism is the position that the best scientific strategy is to attempt to reduce explanations to the smallest possible entities. Methodological reductionism would thus include the claim that the atomic explanation of a substance's boiling point is preferable to the chemical explanation, and that an explanation based on even smaller particles (quarks and leptons, perhaps) would be even better.[citation needed] Methodological reductionism, therefore, is the opinion that all scientific theories either can or should be reduced to a single super~theory through the process of theoretical reduction.

Theory reductionism

Theory reduction is the process by which one theory absorbs another. For example, both Kepler's laws of the motion of the planets and Galileo's theories of motion formulated for terrestrial objects are reducible to Newtonian theories of mechanics because all the explanatory power of the former are contained within the latter. Furthermore, the reduction is considered to be beneficial because Newtonian mechanics is a more general theory—- that is, it explains more events than Galileo's or Kepler's. Theoretical reduction, therefore, is the reduction of one explanation or theory to another—- that is, it is the absorption of one of our ideas about a particular item into another idea.

In science

Reductionist thinking and methods form the basis for many of the well-developed topics of modern science, including much of physics, chemistry and cell biology. Classical mechanics in particular is seen as a reductionist framework, and statistical mechanics can be considered as a reconciliation of macroscopic thermodynamic laws with the reductionist method of explaining macroscopic properties in terms of microscopic components.

In science, reductionism implies that certain topics of study are based on areas that study smaller spatial scales or organizational units. While it is commonly accepted that the foundations of chemistry are based in physics, and molecular biology is based on chemistry, similar statements become controversial when one considers less rigorously defined intellectual pursuits. For example, claims that sociology is based on psychology, or that economics is based on sociology and psychology would be met with reservations. These claims are difficult to substantiate even though there are obvious associations between these topics (for instance, most would agree that psychology can affect and inform economics). The limit of reductionism's usefulness stems from emergent properties of complex systems, which are more common at certain levels of organization. For example, certain aspects of evolutionary psychology and sociobiology are rejected by some who claim that complex systems are inherently irreducible and that a holistic method is needed to understand them.

Some strong reductionists believe that the behavioral sciences should become "genuine" scientific disciplines based on genetic biology, and on the systematic study of culture (see Richard Dawkins's concept of memes). In his book The Blind Watchmaker, Dawkins introduced the term "hierarchical reductionism"[17] to describe the opinion that complex systems can be described with a hierarchy of organizations, each of which is only described in terms of objects one level down in the hierarchy. He provides the example of a computer, which using hierarchical reductionism is explained in terms of the operation of hard drives, processors, and memory, but not on the level of logic gates, or on the even simpler level of electrons in a semiconductor medium.

Others argue that inappropriate use of reductionism limits our understanding of complex systems. In particular, ecologist Robert Ulanowicz says that science must develop techniques to study ways in which larger scales of organization influence smaller ones, and also ways in which feedback loops create structure at a given level, independently of details at a lower level of organization. He advocates (and uses) information theory as a framework to study propensities in natural systems.[18] Ulanowicz attributes these criticisms of reductionism to the philosopher Karl Popper and biologist Robert Rosen.[19]

The idea that phenomena such as emergence and work within the topic of complex systems theory pose limits to reductionism has been advocated by Stuart Kauffman.[20] Emergence is especially relevant when systems exhibit historicity.[21] Emergence is strongly related to nonlinearity.[22] The limits of the application of reductionism are claimed to be especially evident at levels of organization with higher amounts of complexity, including living cells,[23] neural networks, ecosystems, society, and other systems formed from assemblies of large numbers of diverse components linked by multiple feedback loops.[23][24]

Nobel laureate Philip Warren Anderson used the idea that symmetry breaking is an example of an emergent phenomenon in his 1972 Science paper "More is different" to make an argument about the limitations of reductionism.[25] One observation he made was that the sciences can be arranged roughly in a linear hierarchy — particle physics, solid state physics, chemistry, molecular biology, cellular biology, physiology, psychology, social sciences — in that the elementary entities of one science obeys the principles of the science that precedes it in the hierarchy; yet this does not imply that one science is just an applied version of the science that precedes it. He writes that "At each stage, entirely new laws, concepts and generalizations are necessary, requiring inspiration and creativity to just as great a degree as in the previous one. Psychology is not applied biology nor is biology applied chemistry."

Disciplines such as cybernetics and systems theory imply non-reductionism, sometimes to the extent of explaining phenomena at a given level of hierarchy in terms of phenomena at a higher level, in a sense, the opposite of reductionism.[26]

In mathematics

In mathematics, reductionism can be interpreted as the philosophy that all mathematics can (or ought to) be based on a common foundation, which for modern mathematics is usually axiomatic set theory.

Ernst Zermelo was one of the major advocates of such an opinion; he also developed much of axiomatic set theory. It has been argued that the generally accepted method of justifying mathematical axioms by their usefulness in common practice can potentially weaken Zermelo's reductionist claim.[27]

Jouko Väänänen has argued for second-order logic as a foundation for mathematics instead of set theory,[28] whereas others have argued for category theory as a foundation for certain aspects of mathematics.[29][30]

The incompleteness theorems of Kurt Gödel, published during 1931, caused doubt about the attainability of an axiomatic foundation for all of mathematics. Any such foundation would have to include axioms powerful enough to describe the arithmetic of the natural numbers (a subset of all mathematics). Yet Gödel proved that for any self-consistent recursive axiomatic system powerful enough to describe the arithmetic of the natural numbers, there are propositions about the natural numbers that cannot be proved from the axioms, but which we can prove in the natural language with which we described the axioms. Such propositions are known as formally undecidable propositions. For example, the continuum hypothesis is undecidable in the Zermelo-Fraenkel set theory as shown by Cohen.

In religion

Religious reductionism generally attempts to explain religion by explaining it in terms of nonreligious causes. A few examples of reductionistic explanations for the presence of religion are: that religion can be reduced to humanity's conceptions of right and wrong, that religion is fundamentally a primitive attempt at controlling our environments, that religion is a way to explain the existence of a physical world, and that religion confers an enhanced survivability for members of a group and so is reinforced by natural selection.[31] Anthropologists Edward Burnett Tylor and James George Frazer employed some religious reductionist arguments.[32] Sigmund Freud held that religion is nothing more than an illusion, or even a mental illness, and Marx claimed that religion is "the sigh of the oppressed," and the opium of the people providing only "the illusory happiness of the people," thus providing two influential examples of reductionistic views against the idea of religion.

In linguistics

Linguistic reductionism is the idea that everything can be described or explained by a language with a limited number of concepts, and combinations of those concepts.[33] An example is the language Toki Pona.

In philosophy

The concept of downward causation poses an alternative to reductionism within philosophy. This opinion is developed by Peter Bøgh Andersen, Claus Emmeche, Niels Ole Finnemann, and Peder Voetmann Christiansen, among others. These philosophers explore ways in which one can talk about phenomena at a larger-scale level of organization exerting causal influence on a smaller-scale level, and find that some, but not all proposed types of downward causation are compatible with science. In particular, they find that constraint is one way in which downward causation can operate.[34] The notion of causality as constraint has also been explored as a way to shed light on scientific concepts such as self-organization, natural selection, adaptation, and control.[35]

Free will

Philosophers of the Enlightenment worked to insulate human free will from reductionism. Descartes separated the material world of mechanical necessity from the world of mental free will. German philosophers introduced the concept of the "noumenal" realm that is not governed by the deterministic laws of "phenomenal" nature, where every event is completely determined by chains of causality.[36] The most influential formulation was by Immanuel Kant, who distinguished between the causal deterministic framework the mind imposes on the world—- the phenomenal realm—- and the world as it exists for itself, the noumenal realm, which included free will. To insulate theology from reductionism, 19th century post-Enlightenment German theologians, especially Friedrich Schleiermacher and Albrecht Ritschl, used the Romantic method of basing religion on the human spirit, so that it is a person's feeling or sensibility about spiritual matters that comprises religion.[37]

Antireductionism

The anti-reductionist considers as minimum requirement upon the reductionist: "At the very least the anti-reductionist is owed an account of why the intuitions arise if they are not accurate."[38]

A contrast to reductionism is holism or emergentism. Holism is the idea that items can have properties, (emergent properties), as a whole that are not explainable from the sum of their parts. The principle of holism was summarized concisely by Aristotle in the Metaphysics: "The whole is more than the sum of its parts".

Alternatives

The development of systems thinking has provided methods for describing issues in a holistic rather than a reductionist way, and many scientists use a holistic paradigm.[39] When the terms are used in a scientific context, holism and reductionism refer primarily to what sorts of models or theories offer valid explanations of the natural world; the scientific method of falsifying hypotheses, checking empirical data against theory, is largely unchanged, but the method guides which theories are considered. The conflict between reductionism and holism in science is not universal—- it usually concerns whether or not a holistic or reductionist method is appropriate in the context of studying a specific system or phenomenon.

In many cases (such as the kinetic theory of gases), given a good understanding of the components of the system, one can predict all the important properties of the system as a whole. In other systems, emergent properties of the system are said to be almost impossible to predict from knowledge of the parts of the system. Complexity theory studies systems and properties of the latter type.

Alfred North Whitehead's metaphysics opposed reductionism. He refers to this as the "fallacy of the misplaced concreteness". His scheme was to frame a rational, general understanding of phenomena, derived from our reality.

Sven Erik Jorgensen, an ecologist, states both theoretical and practical arguments for a holistic method in certain topics of science, especially ecology. He argues that many systems are so complex that it will not ever be possible to describe all their details. Making an analogy to the Heisenberg uncertainty principle in physics, he argues that many interesting and relevant ecological phenomena cannot be replicated in laboratory conditions, and thus cannot be measured or observed without influencing and changing the system in some way. He also indicates the importance of interconnectedness in biological systems. His opinion is that science can only progress by outlining what questions are unanswerable and by using models that do not attempt to explain everything in terms of smaller hierarchical levels of organization, but instead model them on the scale of the system itself, taking into account some (but not all) factors from levels both higher and lower in the hierarchy.[40]

Criticism

Fragmentalism is an alternative term for ontological reductionism,[41] although fragmentalism is frequently used in a pejorative sense.[42] Anti-realists use the term fragmentalism in arguments that the world does not exist of separable entities, instead consisting of wholes. For example, advocates of this idea claim that:
The linear deterministic approach to nature and technology promoted a fragmented perception of reality, and a loss of the ability to foresee, to adequately evaluate, in all their complexity, global crises in ecology, civilization and education.[43]
The term "fragmentalism" is usually applied to reductionist modes of thought, frequently with the related pejorative term of scientism. This usage is popular amongst some ecological activists:
There is a need now to move away from scientism and the ideology of cause-and-effect determinism toward a radical empiricism, such as William James proposed, as an epistemology of science.[44]
These perspectives are not new and during the early twentieth century, William James noted that rationalist science emphasized what he termed fragmentation and disconnection.[45]

Such opinions also motivate many criticisms of the scientific method:
The scientific method only acknowledges monophasic consciousness. The method is a specialized system that emphasizes studying small and distinctive parts in isolation, which results in fragmented knowledge.[45]
An alternative usage of this term is in cognitive psychology. Here, George Kelly developed "constructive alternativism" as a form of personal construct psychology, this provided an alternative to what he considered "accumulative fragmentalism". For this theory, knowledge is seen as the construction of successful mental models of the exterior world, rather than the accumulation of independent "nuggets of truth".[46]

Equality (mathematics)

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