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Putamen

The putamen (from Latin, meaning "nutshell") is a round structure located at the base of the forebrain (telencephalon). The putamen and caudate nucleus together form the dorsal striatum. It is also one of the structures that compose the basal nuclei. Through various pathways, the putamen is connected to the substantia nigra, the globus pallidus, the claustrum, and the thalamus, in addition to many regions of the cerebral cortex. A primary function of the putamen is to regulate movements at various stages (e.g. preparation and execution) and influence various types of learning. It employs GABA, acetylcholine, and enkephalin to perform its functions. The putamen also plays a role in degenerative neurological disorders, such as Parkinson's disease.

History
The word "putamen" is from Latin, referring to that which "falls off in pruning", from "putare", meaning "to prune, to think, or to consider".

Until recently, most MRI research focused broadly on the basal ganglia as a whole, for various reasons (e.g. image resolution, rarity of isolated infarct or hemorrhage within the putamen, etc.). However, many studies have been done on the basal ganglia and relevant brain-behavior relationships. In the 1970s, the first single unit recordings were done with monkeys monitoring pallidal neuron activity related to movement. Since then, more extensive neuronal tracing, stimulation, and imaging research methods (e.g. fMRI, DWI) that allow for investigation of the putamen have been developed.

Anatomy
The putamen is a structure in the forebrain. Along with the caudate nucleus it forms the dorsal striatum. The caudate and putamen contain the same types of neurons and circuits – many neuroanatomists consider the dorsal striatum to be a single structure, divided into two parts by a large fiber tract, the internal capsule, passing through the middle. The putamen, together with the globus pallidus, makes up the lentiform nucleus. The putamen is the outermost portion of the basal ganglia. These are a group of nuclei in the brain that are interconnected with the cerebral cortex, thalamus, and brainstem. Basal ganglia include the dorsal striatum, substantia nigra, nucleus accumbens, and the subthalamic nucleus.

In mammals, the basal ganglia are associated with motor control, cognition, emotions, learning, and domain-general functions important for executive functioning as well as support for domain-specific languages. The basal ganglia are located bilaterally, and have rostral and caudal divisions. The putamen is located in the rostral division as part of the striatum. The basal ganglia receive input from the cerebral cortex, via the striatum.

The putamen is interconnected with the following structures:

This description is rudimentary and does not nearly exhaust even the basic established circuitry of the putamen. The cortico-subcortico-cortical circuits with putaminal involvement are dense and complicated, consisting of a wide range of axonal, dendritic, chemical, afferent, and efferent substrates. The putamen's outputs are highly arborized across output structures, and cortical efferents arise from layers III-VI of the cortex, dependent on gyri and location within the putamen. Topographical organization of the putamen combines the following elements: anterior-to-posterior functional and somatotopic gradients, lateral-to-medial functional and somatotopic gradients, diffuse terminal output, patchy localized terminal output, segregated terminals from adjacent regions, finely interdigitated terminals from distal cortical regions in a seemingly overlapping fashion.

Caudate nucleus
The caudate works with the putamen to receive the input from cerebral cortex. Collectively, they can be considered the "entrance" to the basal ganglia. Projections from the putamen reach the caudate directly via the caudolenticular grey bridges. The putamen and caudate are jointly connected with the substantia nigra, however the caudate outputs more densely to the substantia nigra pars reticulata while the putamen sends more afferents to the internal globus pallidus.

Substantia nigra
The substantia nigra contains two parts: the substantia nigra pars compacta (SNpc) and the substantia nigra pars reticulata (SNpr). The SNpc obtains input from the putamen and caudate, and sends information back. The SNpr also obtains input from the putamen and caudate. However, it sends the input outside the basal ganglia to control head and eye movements. The SNpc produces dopamine, which is crucial for movements. The SNpc degenerates during Parkinson's disease.

Globus pallidus
The globus pallidus contains two parts: the globus pallidus pars externa (GPe) and the globus pallidus pars interna (GPi). Both regions acquire input from the putamen and caudate and communicate with the subthalamic nucleus. However, mostly the GPi sends GABAergic inhibitory output to the thalamus. The GPi also sends projections to parts of the midbrain, which have been assumed to affect posture control.

Types of pathways
The putamen (and striatum in general) has numerous, parallel circuits that allow for cortico-subcortico-cortico communication loops. These have been described, broadly, as the direct, indirect, and hyper direct pathways. GABAergic projections of the putamen have an inhibitory effect on the thalamus. Thalamic projections from the centromedian and parafascicular nuclei have an excitatory effect on the putamen. Unlike the thalamus, which has broad reciprocal connectivity, cortical projections with the putamen are afferent, thus sending information as opposed to receiving it. Cortical communication is accomplished via multi-fiber pathways as outlined previously (i.e. via other subcortical structures).

Dopamine
Dopamine is a neurotransmitter that has a dominant role in the putamen, most of it is supplied from the substantia nigra. When a cell body of a neuron (in the putamen or caudate nuclei) fires an action potential, dopamine is released from the presynaptic terminal. Since projections from the putamen and caudate nuclei modulate the dendrites of the substantia nigra, the dopamine influences the substantia nigra, which affects motor planning. This same mechanism is involved in drug addiction. In order to control the amount of dopamine in the synaptic cleft, and the amount of dopamine binding to post synaptic terminals, presynaptic dopaminergic neurons function to reuptake the excess dopamine.

Other neurotransmitters
The putamen also plays a role in modulation of other neurotransmitters. It releases GABA, enkephalin, substance P, and acetylcholine. It receives serotonin and glutamate.

Function: motor skills
The putamen is interconnected with many other structures, and works in conjunction with them to influence many types of motor behaviors. These include motor planning, learning, and execution, motor preparation, specifying amplitudes of movement, and movement sequences.

Some neurologists hypothesize that the putamen also plays a role in the selection of movement (e.g. Tourette syndrome) and the "automatic" performance of previously learned movements (e.g. Parkinson's disease).

In one study it was found that the putamen controls limb movement. The goal of this study was to determine whether particular cell activity in the putamen of primates was related to the direction of limb movement or to the underlying pattern of muscular activity. Two monkeys were trained to perform tasks that involved the movement of loads. The tasks were created so that movement could be distinguished from muscle activity. Neurons in the putamen were selected for monitoring only if they were related both to the task and to arm movements outside the task. It was shown that 50% of the neurons that were monitored were related to the direction of movement, independent of the load.

Another study was done to investigate movement extent and speed using PET mapping of regional cerebral blood flow in 13 humans. Movement tasks were performed with a joystick-controlled cursor. Statistical tests were done to calculate the extent of movements and what regions of the brain the movements correlate to. It was found that "increasing movement extent was associated with parallel increases of rCBF in bilateral basal ganglia (BG; putamen and globus pallidus) and ipsilateral cerebellum." This not only shows that the putamen affects movement but it also shows that the putamen integrates with other structures in order to perform tasks.

One study was done in order to specifically investigate how the basal ganglia influences the learning of sequential movements. Two monkeys were trained to press a series of buttons in sequence. The methods used were designed to be able to monitor the well-learned tasks versus the new tasks. Muscimol was injected into various parts of the basal ganglia, and it was found that "the learning of new sequences became deficient after injections in the anterior caudate and putamen, but not the middle-posterior putamen". This shows that different areas of the striatum are utilized when performing various aspects of the learning of sequential movements.

Role in learning
In many studies, it has become apparent that the putamen plays a role in many types of learning. Some examples are listed below:

Reinforcement and implicit learning
Along with various types of movement, the putamen also affects reinforcement learning and implicit learning.

Reinforcement learning is interacting with the environment and catering actions to maximize the outcome. Implicit learning is a passive process where people are exposed to information and acquire knowledge through exposure. Although the exact mechanisms are not known, it is clear that dopamine and tonically active neurons play a key role here. Tonically active neurons are cholinergic interneurons that fire during the entire duration of the stimulus and fire at about 0.5–3 impulses per second. Phasic neurons are the opposite and only fire an action potential when movement occurs.

Category learning
One particular study used patients with focal lesions on the basal ganglia (specifically the putamen) due to stroke in order to study category learning. The advantage to using these types of patients is that dopaminergic projections to the prefrontal cortex are more likely to be intact. Also, in these patients, it is easier to relate specific brain structures to function because the lesion only occurs in a specific place. The goal of this study was to determine whether or not these lesions affect rule-based and information-integration task learning. Rule-based tasks are learned via hypothesis-testing dependent on working memory. Information-integration tasks are ones wherein the accuracy is maximized when information from two sources are integrated at a pre-decisional stage, which follows a procedural-based system.

Seven participants with basal ganglia lesions were used in the experiment, along with nine control participants. It is important to note that the caudate was not affected. The participants were tested for each type of learning during separate sessions, so the information processes would not interfere with each other. During each session, participants sat in front of a computer screen and various lines were displayed. These lines were created by using a randomization technique where random samples were taken from one of four categories. For ruled-based testing, these samples were used to construct lines of various length and orientation that fell into these four separate categories. After the stimulus was displayed, the subjects were asked to press 1 of 4 buttons to indicate which category the line fell into. The same process was repeated for information-integration tasks, and the same stimuli were used, except that the category boundaries were rotated 45°. This rotation causes the subject to integrate the quantitative information about the line before determining what category it is in.

It was found that subjects in the experimental group were impaired while performing rule-based tasks, but not information-integration ones. After statistical testing, it was also hypothesized that the brain began using information-integration techniques to solve the rule-based learning tasks. Since rule-based tasks use the hypothesis-testing system of the brain, it can be concluded that the hypothesis-testing system of the brain was damaged/weakened. It is known that the caudate and working memories are part of this system. Therefore, it was confirmed that the putamen is involved in category learning, competition between the systems, feed-back processing in rule-based tasks, and is involved in the processing of pre-frontal regions (which relate to working memory and executive functioning). Now it is known that not only the basal ganglia and caudate affect category learning.

Role in "hate circuit"
Tentative studies have suggested that the putamen may play a role in the so-called "hate circuit" of the brain. A recent study was done in London by the department of cell and developmental biology at University College London. An fMRI was done on patients while they viewed a picture of people they hated and people who were "neutral". During the experiment, a "hate score" was recorded for each picture. The activity in sub-cortical areas of the brain implied that the "hate circuit" involves the putamen and the insula. It has been theorized that the "putamen plays a role in the perception of contempt and disgust, and may be part of the motor system that's mobilized to take action." It was also found that the amount of activity in the hate circuit correlates with the amount of hate a person declares, which could have legal implications concerning malicious crimes.

Role in transgender individuals
The putamen of transgender women was found to have significantly larger amounts of grey matter compared to the putamen of cisgender men. This possibly suggests that a difference in the putamen may exist between trans women and cisgender men.

Parkinson's disease
After discovering the function of the putamen, it has become apparent to neurologists that the putamen and other parts of the basal ganglia play an important role in Parkinson's disease and other diseases that involve the degeneration of neurons.

Parkinson's disease is the slow and steady loss of dopaminergic neurons in substantia nigra pars compacta. In Parkinson's disease the putamen plays a key role because its inputs and outputs are interconnected to the substantia nigra and the globus pallidus. In Parkinson's disease the activity in direct pathways to interior globus pallidus decreases and activity in indirect pathways to external globus pallidus increases. It has also been noted that Parkinson's patients have a difficult time with motor planning.

Other diseases and disorders
The following diseases and disorders are linked with the putamen:

 * Cognitive decline in Alzheimer's disease
 * Huntington's disease
 * Wilson's disease
 * Dementia with Lewy bodies
 * Corticobasal degeneration
 * Tourette syndrome
 * Schizophrenia
 * Depression
 * Attention deficit hyperactivity disorder
 * Chorea
 * Obsessive-Compulsive Disorder
 * Kernicterus
 * Other anxiety disorders

In other animals
The putamen in humans is relatively similar in structure and function to other animals. Therefore, many studies on the putamen have been done on animals (monkeys, rats, cats, etc.), as well as humans. However, inter-species variation are indeed observed in mammals, and have been documented for white matter putaminal connectivity. Variation is primarily related to structural connectivity patterns, while somatotopic organization principles are retained. Primate research since the 1980s through to the present has established that cortical regions relation to higher-order cognition primarily send afferent neurons to the rostal-most portion of the putamen, while the remainder of this structure in primates primarily serves sensori-motor functions and is densely interconnected with primary and supplementary motor regions.

title: Putamen


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Striatum

The striatum (: striata) or corpus striatum (also called the striate nucleus) is a nucleus (a cluster of neurons) in the subcortical basal ganglia of the forebrain. The striatum is a critical component of the motor and reward systems; receives glutamatergic and dopaminergic inputs from different sources; and serves as the primary input to the rest of the basal ganglia.

Functionally, the striatum coordinates multiple aspects of cognition, including both motor and action planning, decision-making, motivation, reinforcement, and reward perception. The striatum is made up of the caudate nucleus and the lentiform nucleus. However, some authors believe it is made up of caudate nucleus, putamen, and ventral striatum. The lentiform nucleus is made up of the larger putamen, and the smaller globus pallidus. Strictly speaking the globus pallidus is part of the striatum. It is common practice, however, to implicitly exclude the globus pallidus when referring to striatal structures.

In primates, the striatum is divided into the ventral striatum and the dorsal striatum, subdivisions that are based upon function and connections. The ventral striatum consists of the nucleus accumbens and the olfactory tubercle. The dorsal striatum consists of the caudate nucleus and the putamen. A white matter nerve tract (the internal capsule) in the dorsal striatum separates the caudate nucleus and the putamen. Anatomically, the term striatum describes its striped (striated) appearance of grey-and-white matter.

Structure

The striatum is the largest structure of the basal ganglia. The striatum is divided into a ventral subdivision and a dorsal subdivision, based upon function and connections.

The ventral striatum is composed of the nucleus accumbens and the olfactory tubercle. The nucleus accumbens is made up of the nucleus accumbens core and the nucleus accumbens shell, which differ by neural populations. The olfactory tubercle receives input from the olfactory bulb but has not been shown to play a role in processing smell. In non-primate species, the islands of Calleja are included. The ventral striatum is associated with the limbic system and has been implicated as a vital part of the circuitry for decision making and reward-related behavior.

The dorsal striatum is composed of the caudate nucleus and the putamen.

Staining can differentiate the striatum into two distinct compartments of striosomes or patches, and a surrounding matrix; this is particularly evident on the components of acetylcholinesterase and calbindin. More studies have been carried out on the dorsal striatum but the compartments have also been identified in the ventral striatum. In the dorsal striatum, striosomes make up 10-15% of the striatal volume.

Cell types
Types of cells in the striatum include:
 * Medium spiny neurons (MSNs), which are the principal neurons of the striatum. They are GABAergic and, thus, are classified as inhibitory neurons. Medium spiny projection neurons comprise 95% of the total neuronal population of the human striatum. Medium spiny neurons have two characteristic types: D1-type MSNs and D2-type MSNs.  A subpopulation of MSNs contain both D1-type and D2-type receptors, with approximately 40% of striatal MSNs expressing both DRD1 and DRD2 mRNA.
 * Cholinergic interneurons release acetylcholine, which has a variety of important effects in the striatum. In humans, other primates, and rodents, these interneurons respond to salient environmental stimuli with stereotyped responses that are temporally aligned with the responses of dopaminergic neurons of the substantia nigra. The large aspiny cholinergic interneurons themselves are affected by dopamine through D5 dopamine receptors. Dopamine also directly controls communication between cholinergic interneurons.
 * There are many types of GABAergic interneurons. The best known are parvalbumin expressing interneurons, also known as fast-spiking interneurons, which participate in powerful feedforward inhibition of principal neurons. Also, there are GABAergic interneurons that express tyrosine hydroxylase, somatostatin, nitric oxide synthase and neuropeptide-y. Recently, two types of neuropeptide-y expressing GABAergic interneurons have been described in detail, one of which translates synchronous activity of cholinergic interneurons into inhibition of principal neurons. These neurons of the striatum are not distributed evenly.

There are two regions of neurogenesis in the brain – the subventricular zone (SVZ) in the lateral ventricles, and the dentate gyrus in the hippocampal formation. Neuroblasts that form in the lateral ventricle adjacent to the striatum, integrate in the striatum. This has been noted in the human striatum following an ischemic stroke. Injury caused to the striatum stimulates the migration of neuroblasts from the SVZ, to the striatum, where they differentiate into adult neurons. The normal passage of SVZ neuroblasts is to the olfactory bulb but this traffic is diverted to the striatum after an ischemic stroke. However, few of the new developed neurons survive.

Inputs
The largest connection is from the cortex, in terms of cell axons. Many parts of the neocortex innervate the dorsal striatum. The cortical pyramidal neurons projecting to the striatum are located in layers II-VI, with the most dense projections come from layer V.  They end mainly on the dendritic spines of the spiny neurons. They are glutamatergic, exciting striatal neurons.

The striatum is seen as having its own internal microcircuitry. The ventral striatum receives direct input from multiple regions in the cerebral cortex and limbic structures such as the amygdala, thalamus, and hippocampus, as well as the entorhinal cortex and the inferior temporal gyrus. Its primary input is to the basal ganglia system. Additionally, the mesolimbic pathway projects from the ventral tegmental area to the nucleus accumbens of the ventral striatum.

Another well-known afferent is the nigrostriatal connection arising from the neurons of the substantia nigra pars compacta. While cortical axons synapse mainly on spine heads of spiny neurons, nigral axons synapse mainly on spine shafts. In primates, the thalamostriatal afferent comes from the central median-parafascicular complex of the thalamus (see primate basal ganglia system). This afferent is glutamatergic. The participation of truly intralaminar neurons is much more limited. The striatum also receives afferents from other elements of the basal ganglia such as the subthalamic nucleus (glutamatergic) or the external globus pallidus (GABAergic).

Targets
The primary outputs of the ventral striatum project to the ventral pallidum, then the medial dorsal nucleus of the thalamus, which is part of the frontostriatal circuit. Additionally, the ventral striatum projects to the globus pallidus, and substantia nigra pars reticulata. Some of its other outputs include projections to the extended amygdala, lateral hypothalamus, and pedunculopontine nucleus.

Striatal outputs from both the dorsal and ventral components are primarily composed of medium spiny neurons (MSNs), a type of projection neuron, which have two primary phenotypes: "indirect" MSNs that express D2-like receptors and "direct" MSNs that express D1-like receptors.

The main nucleus of the basal ganglia is the striatum which projects directly to the globus pallidus via a pathway of striatopallidal fibers. The striato-pallidal pathway has a whitish appearance due to the myelinated fibers. This projection comprises successively the external globus pallidus (GPe), the internal globus pallidus (GPi), the pars compacta of the substantia nigra (SNc), and the pars reticulata of substantia nigra (SNr). The neurons of this projection are inhibited by GABAergic synapses from the dorsal striatum. Among these targets, the GPe does not send axons outside the system. Others send axons to the superior colliculus. Two others comprise the output to the thalamus, forming two separate channels: one through the internal segment of the globus pallidus to the ventral oralis nuclei of the thalamus and from there to the cortical supplementary motor area and another through the substantia nigra to the ventral anterior nuclei of the thalamus and from there to the frontal cortex and the occulomotor cortex.

Blood supply
Deep penetrating striate arteries supply blood to the striatum. These arteries include the recurrent artery of Heubner arising from the anterior cerebral artery, and the lenticulostriate arteries arising from the middle cerebral artery.

Function
The ventral striatum, and the nucleus accumbens in particular, primarily mediates reward, cognition, reinforcement, and motivational salience. By contrast, the dorsal striatum primarily mediates cognition involving motor function, certain executive functions (e.g., inhibitory control and impulsivity), and stimulus-response learning. There is a small degree of overlap, as the dorsal striatum is also a component of the reward system that, along with the nucleus accumbens core, mediates the encoding of new motor programs associated with future reward acquisition (e.g., the conditioned motor response to a reward cue).

The striatum is also thought to play a role in an at least partially dissociable executive control network for language, applied to both verbal working memory and verbal attention. These models take the form of a frontal-striatal network for language processing. While the striatum is often not included in models of language processing, as most models only include cortical regions, integrative models are becoming more popular in light of imaging studies, lesion studies on aphasic patients, and studies of language disorders concomitant with diseases known to affect the striatum like Parkinson's and Huntington's disease.

Metabotropic dopamine receptors are present both on spiny neurons and on cortical axon terminals. Second messenger cascades triggered by activation of these dopamine receptors can modulate pre- and postsynaptic function, both in the short term and in the long term. In humans, the striatum is activated by stimuli associated with reward, but also by aversive, novel, unexpected, or intense stimuli, and cues associated with such events. fMRI evidence suggests that the common property linking these stimuli, to which the striatum is reacting, is salience under the conditions of presentation. A number of other brain areas and circuits are also related to reward, such as frontal areas. Functional maps of the striatum reveal interactions with widely distributed regions of the cerebral cortex important to a diverse range of functions.

The interplay between the striatum and the prefrontal cortex is relevant for behavior, particularly adolescent development as proposed by the dual systems model.

Parkinson's disease and other movement disorders
Parkinson's disease results in loss of dopaminergic innervation to the dorsal striatum (and other basal ganglia) and a cascade of consequences. Atrophy of the striatum is also involved in Huntington's disease, and movement disorders such as chorea, choreoathetosis, and dyskinesias. These have also been described as circuit disorders of the basal ganglia.

Addiction
Addiction, a disorder of the brain's reward system, arises through the overexpression of DeltaFosB (ΔFosB), a transcription factor, in the D1-type medium spiny neurons of the ventral striatum. ΔFosB is an inducible gene which is increasingly expressed in the nucleus accumbens as a result of repeatedly using an addictive drug or overexposure to other addictive stimuli.

Bipolar disorder
An association has been observed between striatal expression of variants of the PDE10A gene and some bipolar I disorder patients. Variants of other genes, DISC1 and GNAS, have been associated with bipolar II disorder.

Autism spectrum disorder
Autism spectrum disorder (ASD) is characterized by cognitive inflexibility and poor understanding of social systems. This inflexible behavior originates in defects in the pre-frontal cortex as well as the striatal circuits. The defects in the striatum seem to specifically contribute to the motor, social and communication impairments seen in ASD patients. In mice which have an ASD-like phenotype induced via the overexpression of the eukaryotic initiation of translation factor 4E, it has been shown that these defects seem to stem from the reduced ability to store and process information in the striatum, which leads to the difficulty seen in forming new motor patterns, as well as disengaging from existing ones.

Dysfunction
Dysfunction in the ventral striatum can lead to a variety of disorders, most notably depression and obsessive-compulsive disorder. Because of its involvement in reward pathways, the ventral striatum has also been implicated in playing a critical role in addiction. It has been well established that the ventral striatum is strongly involved in mediating the reinforcing effects of drugs, especially stimulants, through dopaminergic stimulation.

Language disorders
Lesions to the striatum have been associated with deficits in speech production and comprehension. While striatal damage can impact all levels of language, damage can broadly be characterized as affecting the ability to manipulate linguistic units and rules, resulting in the promotion of default linguistic forms in conflicting situations in which selection, inhibition, and monitoring load is increased. Two subregions of the striatum have been shown to be particularly important in language: the caudate nucleus and left putamen. Lesions localized to the caudate nucleus, as well as direct electrical stimulation, can result in lexical paraphasias and perservations (continuations of an utterance after the stimulus has ceased), which is associated with inhibited executive control, in the sense that executive control allows for the selection of the best choice among competing alternatives). Stimulation of the putamen results in the inhibition of articulatory sequences and the inability to initiate motor speech commands.

History
In the seventeenth and eighteenth centuries, the term corpus striatum was used to designate many distinct, deep, infracortical elements of the hemisphere. Etymologically, it is derived from (Latin) striatus = "grooved, striated" and the English striated = having parallel lines or grooves on the surface. In 1876 David Ferrier contributed decades of research to the subject; concluding that the corpus striatum was vital in the "organization and generation of voluntary movement". In 1941, Cécile and Oskar Vogt simplified the nomenclature by proposing the term striatum for all elements in the basal ganglia built with striatal elements: the caudate nucleus, the putamen, and the fundus striati, which is the ventral part linking the two preceding together ventrally to the inferior part of the internal capsule.

The term neostriatum was coined by comparative anatomists comparing the subcortical structures between vertebrates, because it was thought to be a phylogenetically newer section of the corpus striatum. The term is still used by some sources, including Medical Subject Headings.

Other animals
In birds the term used was the paleostriatum augmentatum, and in the new avian terminology listing (as of 2002) for neostriatum this has been changed to the nidopallium.

In non-primate species, the islands of Calleja are included in the ventral striatum.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Striatum


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Voicelessness

In linguistics, voicelessness is the property of sounds being pronounced without the larynx vibrating. Phonologically, it is a type of phonation, which contrasts with other states of the larynx, but some object that the word phonation implies voicing and that voicelessness is the lack of phonation.

The International Phonetic Alphabet (IPA) has distinct letters for many voiceless and modally voiced pairs of consonants (the obstruents), such as. Also, there are diacritics for voicelessness,  and, which is used for letters with a descender. Diacritics are typically used with letters for prototypically voiced sounds, such as vowels and sonorant consonants:. In Russian use of the IPA, the voicing diacritic may be turned for voicelessness, e.g. ⟨ṋ⟩.

Voiceless vowels and other sonorants
Sonorants are sounds such as vowels and nasals that are voiced in most of the world's languages. However, in some languages sonorants may be voiceless, usually allophonically. For example, the Japanese word sukiyaki is pronounced and may sound like  to an English speaker, but the lips can be seen to compress for the. Something similar happens in English words like peculiar and potato.

Voiceless vowels are also an areal feature in languages of the American Southwest (like Hopi and Keres), the Great Basin (including all Numic languages), and the Great Plains, where they are present in Numic Comanche but also in Algonquian Cheyenne, and the Caddoan language Arikara. It also occurs in Woleaian, in contrast to the other Micronesian languages, which instead delete it outright.

Sonorants may also be contrastively, not just environmentally, voiceless. Standard Tibetan, for example, has a voiceless in Lhasa, which sounds similar to but is less noisy than the voiceless lateral fricative  in Welsh; it contrasts with a modally voiced. Welsh contrasts several voiceless sonorants:, , , and , the last represented by "rh".

In Moksha, there is even a voiceless palatal approximant (written in Cyrillic as $⟨йх⟩$ jh) along with  and  (written as $⟨лх⟩$ lh and $⟨рх⟩$ rh). The last two have palatalized counterparts and  ($⟨льх⟩$ and $⟨рьх⟩$). Kildin Sami has also $⟨ҋ⟩$.

Contrastively voiceless vowels have been reported several times without ever being verified (L&M 1996:315).

Lack of voicing contrast in obstruents
Many languages lack a distinction between voiced and voiceless obstruents (stops, affricates, and fricatives). This is the case in nearly all Australian languages, and is widespread elsewhere, for example in Mandarin Chinese, Korean, Danish, Estonian and the Polynesian languages.

In many such languages, obstruents are realized as voiced in voiced environments, such as between vowels or between a vowel and a nasal, and voiceless elsewhere, such as at the beginning or end of the word or next to another obstruent. That is the case in Dravidian and Australian languages and in Korean but not in Mandarin or Polynesian. Usually, the variable sounds are transcribed with the voiceless IPA letters, but for Australian languages, the letters for voiced consonants are often used.

It appears that voicelessness is not a single phenomenon in such languages. In some, such as the Polynesian languages, the vocal folds are required to actively open to allow an unimpeded (silent) airstream, which is sometimes called a breathed phonation (not to be confused with breathy voice). In others, such as many Australian languages, voicing ceases during the hold of a stop (few Australian languages have any other kind of obstruent) because airflow is insufficient to sustain it, and if the vocal folds open, that is only from passive relaxation.

Thus, Polynesian stops are reported to be held for longer than Australian stops and are seldom voiced, but Australian stops are prone to having voiced variants (L&M 1996:53), and the languages are often represented as having no phonemically voiceless consonants at all.

In Southeast Asia, when stops occur at the end of a word, they are voiceless because the glottis is closed, not open, so they are said to be unphonated (have no phonation) by some phoneticians, who considered "breathed" voicelessness to be a phonation.

Yidiny consonants, with no underlyingly voiceless consonants, are posited.

title: Voicelessness


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BBC News (international TV channel)

BBC News (known as BBC World News until 2023) is an international English-language pay television channel owned by BBC Global News Ltd. – a subsidiary of BBC Studios – and operated by the BBC News division of the BBC. The network carries news bulletins, documentaries, and other factual programmes; its programming is based out of studios in London, Washington, D.C., and Singapore. As of April 2023, the channel largely operates as an international feed of the BBC News channel in the UK, sharing the majority of its schedule.

Launched on 11 March 1991 as BBC World Service Television outside Europe, its name was changed to BBC World on 16 January 1995 and to BBC World News on 21 April 2008 and again to BBC News (International) on 3 April 2023 after its consolidation with the domestic BBC News Channel. According to the BBC, the combined seven channels of the Global News operations have the largest audience market share among all its rivals, with an estimated 99 million viewers weekly in 2016–2017, part of the estimated 121 million weekly audience of all its operations.

Unlike the BBC's domestic channels, it is funded by subscription and advertising revenues, and not by the United Kingdom television licence. As such, the channel is not broadcast in the UK directly, although selected programmes and bulletins have been carried on the domestic BBC News channel (especially during overnight hours), and vice versa (including domestic programmes such as Click and HARDtalk, and during breaking news and special events in the UK).

In April 2023, the BBC began to further consolidate the programming and talent of the two channels as part of a corporation-wide streamlining of operations, with both channels now using the BBC News branding. The international feed remains an advertising-supported service, but the two services are structured to use a common schedule with domestic opt-outs for UK-specific news coverage and programmes.

History
The channel originally started as BBC World Service Television and was a commercial operation. The British government refused to fund the new television service using grant-in-aid. (BBC World Service radio was funded by a grant-in-aid from the Foreign, Commonwealth and Development Office until 2014.) The channel started broadcasting on 11 March 1991, after two weeks of real-time pilots, initially as a half-hour bulletin once a day at 19:00GMT.

On Thursday, 26 January 1995 at 19:00 GMT, BBC World Service Television was split into two services:
 * BBC World started broadcasting on Monday, 16 January 1995 at 19:00 GMT and became a 24-hour English free-to-air international news channel.
 * BBC Prime started broadcasting on Monday, 30 January 1995 at 19:00 GMT and became the BBC's light entertainment channel, later renamed BBC Entertainment.

BBC World's on-air design was changed significantly on 3 April 2000, bringing it closer to the look of its sister channel in the UK which was then known as BBC News 24, the on-air look of which had been redesigned in 1999. The look of both channels was made up of red and cream and designed by Lambie-Nairn, with music based on a style described as 'drums and beeps' composed by David Lowe, a departure from the general orchestral nature of music used by other news programmes.

On 8 December 2003, a second makeover, using the same 'drums and beeps' style music but new graphics took place, although on a much smaller scale than that of 2000. The music was changed slightly, while the main colour scheme became black and red, with studios using frosted glass and white and red colours. Later in 2004, the channel's slogan became Putting News First, replacing Demand a Broader View.

On 21 April 2008, as part of a £550,000 rebranding of the BBC's overall news output and visual identity, the channel was rebranded as BBC World News. It later moved to the renovated studio vacated by BBC News 24 (now the BBC News Channel). New graphics were produced by the Lambie-Nairn agency and the music was reworked by David Lowe.

Move to Broadcasting House
The channel relocated to Broadcasting House from its previous home at Television Centre on 14 January 2013. This was part of the move of BBC News and other audio and vision departments of the BBC into one building in Central London. Broadcasting House was refurbished for £1 billion. A new newsroom and several state-of-the-art studios were built.

Consolidation with the domestic BBC News channel
On 26 May 2022, as part of planned cuts and streamlining across the broadcaster, the BBC announced plans for consolidation of the domestic BBC News channel in the UK with BBC World News. The domestic and international versions would share a larger amount of content while maintaining the ability for opt-outs when necessary. The BBC promoted the service as offering "new flagship programmes built around high-profile journalists, and programmes commissioned for multiple platforms". In preparation for the expansion, the BBC began to add additional staff to its Washington, D.C. bureau.

The first stage of these changes took effect on 3 April 2023, when the BBC World News channel changed its name to BBC News, and began to consolidate its on-air lineups and programming with the UK BBC News channel. This resulted in layoffs of about 50 employees, including presenters David Eades, Joanna Gosling, and Tim Willcox. Despite the changes, the international feed of BBC News – which continues to be distributed by BBC Studios – still opts out of programmes that are specific to UK audiences, such as simulcasts of BBC One news bulletins and Newsnight. Deadline Hollywood has reported that in response to concerns over the arrangement by the telecom and broadcasting regulator Ofcom, BBC News was considering allowing for more UK-specific opt-outs.

Broadcasting
Live news output originates from studios B, C and E in Broadcasting House with some recorded programming from Broadcasting House studios A and D and the BBC Millbank studio, as well as Manchester, Singapore and Washington, D.C. The BBC News newsroom is now part of the new consolidated BBC Newsroom in Broadcasting House along with BBC World Service and UK domestic news services. Previously, the channel was broadcast in 4:3, with the news output fitted into a 14:9 frame for both digital and analogue broadcasting, resulting in black bands at the top and bottom of the screen. On 13 January 2009 at 09:57 GMT, the then-named BBC World News switched its broadcast to 16:9 format, initially in Europe on the Astra 1L satellite, and then, from 20 January 2009, on the Eutelsat Hot Bird 6 satellite which fed other broadcast feeds in the Asian region. The channel ceased broadcasting on analogue satellite on 18 April 2006.

High-definition
As a result of the move to Broadcasting House, BBC News gained new studios and equipment to be able to broadcast in high-definition. On 5 August 2013, the international feed of BBC News was offered as a High Definition (HD) feed across the Middle East when it launched its international HD channel on the Arab Satellite Communications Organization. Arabsat was the BBC's first distribution partner in the Middle East to offer the channel in HD. On 1 April 2015, the then-named BBC World News Channel started broadcasting in high definition on the 11.229 GHz/V transponder on Astra 1KR at the 19.2°E orbital position, available free-to-air to viewers with 60 cm dishes across Europe and coastal North Africa.

Worldwide
BBC News claims to be watched by a weekly audience of 74 million in over 200 countries and territories worldwide. BBC News is most commonly watched as a free-to-air (FTA) channel. The channel is available in Europe and many parts of the world via subscription television providers in cable, satellite, IPTV and streaming platforms.

In the United States, the channel is available through providers such as Cablevision, Comcast, Spectrum, Verizon Fios, and U-verse TV. As of 2023, the American distribution and advertising sales for the channel are handled by AMC Networks, who are the minority partner for the BBC's entertainment channel BBC America.

In addition, BBC News syndicates its daytime and evening news programmes to public television stations throughout the US, originally maintaining a distribution partnership with Garden City, New York-based WLIW that lasted from 1998 until October 2008, when the BBC and WLIW mutually decided not to renew the contract. BBC News subsequently entered into an agreement with Community Television of Southern California, Inc., in which Los Angeles PBS member station KCET (which was a public independent station from 2011 to 2018) would take over distribution rights to BBC World News America (the KCET agreement has since been extended to encompass a half-hour simulcast of the 90-minute-long midday news bulletin GMT, which airs in the US as a morning show, and a weekly edition of the BBC news-magazine Newsnight).

Since June 2019, the distribution of the programme has been handled by Washington, D.C. PBS member WETA, which also produces other networked news and public affairs programmes such as the PBS NewsHour and Washington Week. PBS separately began distributing another programme aired by the channel, Beyond 100 Days, as a tape-delayed late night broadcast on 2 January 2018, as an interim replacement for Charlie Rose. Unlike GMT and BBC World News America, Beyond 100 Days is distributed exclusively to PBS member stations as part of the service's base schedule.

China banned BBC News in 2021, although access had already been heavily restricted before then and what was carried was censored, with parts of the show being blacked out by censors operating live. It was banned because of its coverage of the persecution of Uyghur people in China and in retaliation for CGTN being banned from the British market for violating national broadcast regulations.

Online
The channel is available in the US as part of Sling's World News add-on package.

The international feed of BBC News was available on LiveStation from 2012 until the platform closed in 2016, along with the UK simulcast.

The channel was added to various free ad-supported streaming television (FAST) platforms in the United States, including Pluto TV, Xumo Play, Samsung TV Plus, and Sling Freestream, in March 2024. Apart from advertising, the FAST version of the channel is identical to the version already available through traditional U.S. cable and satellite providers.

United Kingdom
The TV platforms in the UK (i.e. Freeview, Sky, BT TV, Freesat, Virgin) do not officially offer the international feed of BBC News as a standalone full-time channel because it carries, and is funded by, advertising. (The BBC's domestic channels are funded by a television licence fee which households and establishments that want to watch television programmes as they are being broadcast must pay). However, it can be easily received due to its free-to-air status on many European satellite systems, including Astra and Hot Bird, and is available in selected London hotels. The UK's TV platforms, however, do provide the UK feed of BBC News channel, which carries much of the same programming. The international feed of BBC News can also be viewed in the public areas of Broadcasting House (in the lobby and café).

However, some programmes are officially available to UK audiences through the channel's UK feed. Such programmes air on the BBC's domestic channels, and some are available on demand on the BBC's iPlayer. From 00:00 to 05:00 UK time, the top-of-the-hour news bulletins are broadcast on both the UK and international feeds of the channel. At 01:30 on weekdays, Asia Business Report and Sport Today also air on both channels. There is a simulcast of the 05:00 UK edition of The Briefing and Business Briefing on BBC One and the BBC News channel. This programme was previously branded as The World Today (later a generic BBC World News bulletin) and World Business Report respectively. At 08:30 UK time, Worklife airs on the BBC News Channel. BBC News also produces a version of Outside Source at 21:00 UK time Monday–Thursday (shown on the BBC News Channel), World News Today at 19:00 Monday–Friday (shown on BBC Four), and 21:00 Friday–Sunday (shown on the BBC News Channel). World News Today replaced The World, which had been broadcast as a simulcast on BBC Four between 2002 and 2007.

The COVID-19 pandemic saw an increase of simulcasts between BBC News and BBC World News with simulcasting running through the morning (10:00 to 11:00) that was also shared with BBC Two, and the evening (19:00 to 20:00 & 12:00 to 22:00). The additional simulcasting was made permanent in August 2020. Consequently, the two channels now simulcast every day from 10:00 to 11:00; on weekdays from 19:00 to 06:00, apart from the BBC News at Ten and for an hour at 20:00; and between 21:00 to 06:00, apart from the evening BBC One bulletin, over the weekend.

The BBC implemented a wider consolidation of the networks' lineups in April 2023.

Programming
Live news programmes:
 * BBC News – International news.
 * BBC World News America – News from the United States and around the world, live from the BBC's Washington DC bureau.
 * The Context with Christian Fraser in London reports on international news, with a focus on the UK, Europe and the US (previously called 100 Days, 100 Days +, Beyond 100 Days and BBC News with Katy and Christian).
 * Focus on Africa – BBC News's flagship African news programme, with news, business and sport from the continent.
 * Newsday – Live from Singapore, news and analysis from both an Asian and global perspective.
 * Sunday with Laura Kuenssberg – Interviews and analysis of the week's big news from the UK and around the world, including interviews with key politicians and personalities from all walks of life.
 * Verified Live – Analysing and fact-checking the truth behind the news based on available data and facts.
 * BBC News Now – fast-moving breaking news with live interviews and reactions.
 * The World Today – The best of the BBC's global journalism. As well as interviews with leading figures from the arts, culture and entertainment.

Live business and sports programmes: Pre-recorded programmes:
 * BBC Sportsday – International sports news and results.
 * BBC Business Today – The latest business news.
 * Click – Technology programme aimed at non-technical audiences.
 * HARDtalk – In-depth interviews with personalities from all walks of life.
 * Newsnight – Weekly highlights from the daily domestic news programme.
 * Our World – Documentaries.
 * Panorama – Current affairs programme, featuring interviews and investigative reports on a wide variety of subjects.
 *  Reporters – Weekly reports from the BBC's correspondents.
 * Talking Movies
 * The Travel Show

Former programmes:
 * The Briefing – Sally Bundock with news, business, and sport.
 * Dateline London – Foreign correspondents based in London discussing the week's international news.
 * Global – International news and analysis.
 * Live – International news, including business and sport.
 * Impact – Global news as it affected audiences in Asia.
 * Outside Source – Discussion and analysis of news topics.
 * Worklife
 * World News Today – A daily news programme with in-depth analysis. Focus on the UK, Europe, the Middle East and Africa. Presenters included Nancy Kacungira, Karin Giannone, Kasia Madera, Alpa Patel and Philippa Thomas.
 * Asia Business Report – Live from Singapore.
 * World Business Report

BBC News bulletins
Half-hour BBC News bulletins are made available to PBS stations in the US through Los Angeles's KCET, a non-commercial independent public television station which has been separate from PBS since the beginning of 2011 due to a rights-fee dispute (it returned to being a minor PBS member station in 2019 after a merger with the major PBS member-station in the market). 80 to 90% of Americans can receive the bulletins, with PBS member stations having scheduling discretion. The programme is broadcast on several PBS stations in markets such as New York City and Washington DC.

On PBS stations, BBC News is not broadcast with traditional commercials (the breaks are filled with news stories) but omits the Met Office international weather forecast at the end of the programme, replacing it with underwriting announcements. The PBS broadcasts are tape-delayed on some stations.

BBC America formerly aired a three-hour block of BBC News programmes from 05:00 to 08:00 on weekdays, until the stabilisation of the network's carriage in the United States. Met Office forecasts were removed, and it was broadcast with advertisements.

Many airlines around the world also play pre-recorded extracts of BBC News, have text headlines from it, or have a full bulletin available on the in-flight entertainment systems.

Previous bulletins
Another BBC World News programme, the hour-long BBC World News America, aired on BBC America at 19:00 ET. A second broadcast at 22:00 ET ended in 2010 when BBC America introduced a second feed for the western time zones of the US. On 18 February 2011, it was announced that BBC World News America would no longer be broadcast on BBC America and would instead be broadcast only on BBC World and local PBS stations in the US as a 30-minute programme.

The channel also produced short bulletins for public transport services in Singapore and Hong Kong:
 * Singapore Mass Rapid Transit service from Marina Bay to Changi International Airport
 * Hong Kong MTR service from Chek Lap Kok International Airport-Disneyland Resort station to Disneyland Resort line

These broadcasts began with the statement: "Welcome to BBC World News on board the Singapore Mass Rapid Transit and Hong Kong MTR". The short bulletin was updated twice a day and has since been replaced by a similar programme from Mediacorp's CNA.

Travellers on the Heathrow Express rail service between London Paddington and Heathrow Airport could watch a summary of the headlines from BBC World News on the LCD screens provided.

Former presenters

 * Samira Ahmed
 * Karen Bowerman
 * Tony Campion
 * Jonathan Charles
 * Stephen Cole
 * James Dagwell
 * Dharshini David
 * Martine Dennis
 * Juliet Dunlop
 * Maya Even
 * Adrian Finighan
 * Rico Hizon
 * Donald MacCormick
 * Anita McNaught
 * Richard Quest
 * Daniela Ritorto
 * Owen Thomas
 * Alastair Yates

Presentation
BBC News is, for the most part, the same channel around the world; the commercials are intended to be the only differences. However, there are some regional programming variations. For example, several programmes are made exclusively for regional viewings, such as the Indian feeds, and The Record Europe, which is only broadcast in Europe. Also, the weather forecasts focus more on the area the viewer is watching from.

On most feeds of BBC News, when no commercials are being inserted by the cable or satellite provider similar to other channels, the break filler shows promotions for upcoming programmes on the channel. During BBC News, a news story that has not been promoted airs during what would be the television advertisement. This is the case on the broadband versions of BBC News, and on versions of BBC News aired in the US on PBS stations. However, there are some global commercials and sponsorships which air throughout the network.

On 11 September 2007, the break filler was redesigned and now more closely resembles previous versions. The promotional videos now fill the entire screen and are interspersed with news and market updates, schedules, and other information. There is also no longer a unifying music composition. Instead, each 20-second promotional video uses music selected from a handful of themes, which have some unifying musical characteristics. The information screens, such as the 10-second plug for the website or the YouTube channel, and the 15-second weather/time/coming-up screens each feature their theme. The colour theme was updated following the relaunch of the channel in April 2008.

Since its inception, and more so since its extensive association with the BBC News channel, the countdown to the hourly news bulletin has been a feature of the channel's presentation, accompanied by music composed by David Lowe. The current style of countdown features reporters and technical staff in many different locations working to bring news stories to air. The new countdown keeps the same music, but with Chameleon-style branding and a clock relocated to the middle from the bottom right corner, similar in style to that of BBC Alba. The countdown can range from 45 seconds to as little as 3 seconds.

Awards
The then-called BBC World News was named Best International News Channel at the Association for International Broadcasting Awards in November 2006. It won a Peabody Award in 2007 for White Horse Village and another in 2009 for Where Giving Life is a Death Sentence.

title: BBC News (international TV channel)


text:
Talk:Weregild

Untitled
I guess every culture had similar custom: Slavs had "glowczyzna" (glowa == head) which was much the same. Should that be reflected in article? szopen

I've mentioned and linked it. Now all it needs is an article on glowczyzna :-). sjc

I don't think German Wehr is related. I'd guess that this is related to Eng. war, cognate to Romance guerra, etc.--Fitzaubrey 04:30, 28 July 2006 (UTC)

 * I agree - the "were" more closely corresponds with "wer", or 'who' in modern German, and geld orginally meant "worth", not "money". In modern German you have the phrase "er gilt" which means "he is worth" or "he counts for" that I think is more closely related. Elle121 02:34, 2 December 2006 (UTC)

You might want to actually go in to detail as to what "weregild" is exactly, not just cultures who used the system, and it's etymology. Some might not understand by what is written.

 * While I understand that women had a weregild it seems to me like the etymology meant "male person." Were didn't mean man as in person, because they had the word "man." It meant male. Weremen were male persons and wyfmen were female persons, if I remember rightly.75.73.32.46 (talk) 16:22, 22 March 2010 (UTC)

 * Similar payments are required under Sharia law. Pustelnik (talk) 01:00, 17 December 2008 (UTC)

it appears this article was entirely written based on guessing how it probably would have been. I have tried to clean up the most blatant falsehoods, but there remain some highly dubious items. It is plainly wrong that no allowance was made for self-defense, as will be clear to anyone looking at the Anglo-Saxon law codes. The claim that no distinction was made between murder and mere manslaughter may be roughly correct, but I suspect that there are exceptions even there. The claim that women  commanded higher amounts, was true for some codes and not for others. The fanciful "explanation" that women were more precious because the Northern Europeans "were nomads" was just pulled out of thin air (aka WP:OR). --dab (𒁳) 21:00, 15 April 2009 (UTC)

How come this article has the "Anglo-Saxon paganism and mythology" template at the bottom? The concept of weregild is by no means paganism or mythology, it's legal history. —Preceding unsigned comment added by 68.167.61.134 (talk) 11:17, 20 July 2010 (UTC)

Removed recent format-breaking changes with value-laden and misplaced editorial comments from 75.52.186.148. Some other forum is probably better suited to signal disapproval of ancient legal practices. — Preceding unsigned comment added by 66.65.140.210 (talk) 05:37, 8 September 2011 (UTC)

Removed some phrases and words from the description following the link to the article on "social hierarchy" because they contained moral stances. More in general, I feel that the "see also" section could be pruned, it contains a lot of links. Gerard RvE (talk) 12:30, 21 June 2012 (UTC)

Is there a reason why Atonement under See Also links specifically to Day of Atonement (Nation of Islam)?? PICURMN (talk) 04:33, 28 July 2015 (UTC)

Rename
According to modern German, money is "Geld" and not "Gild". Shouldn't this article be renamed as Weregeld? Gauravjuvekar (talk) 14:05, 27 December 2011 (UTC)

What was the value of wergild in practical terms?
If a solidus is 4.5g of gold, then a wergild of 200S works out as £28800 today, which is a little over the current average UK wage. Was the contemporary value comparable? (I.e. was the wergild for a common man roughly equivalent to the average annual earnings of the same)? I think this would be a useful statistic to include, if its known Iapetus (talk) 10:18, 7 October 2016 (UTC)

title: Talk:Weregild


text:
Finger

A finger is a prominent digit on the forelimbs of most tetrapod vertebrate animals, especially those with prehensile extremities (i.e. hands) such as humans and other primates. Most tetrapods have five digits (pentadactyly), and short digits (i.e. significantly shorter than the metacarpal/metatarsals) are typically referred to as toes, while those that are notably elongated are called fingers. In humans, the fingers are flexibly articulated and opposable, serving as an important organ of tactile sensation and fine movements, which are crucial to the dexterity of the hands and the ability to grasp and manipulate objects.

Land vertebrate fingers
As terrestrial vertebrates were evolved from lobe-finned fish, their forelimbs are phylogenetically equivalent to the pectoral fins of fish. Within the taxa of the terrestrial vertebrates, the basic pentadactyl plan, and thus also the metacarpals and phalanges, undergo many variations. Morphologically the different fingers of terrestrial vertebrates are homolog. The wings of birds and those of bats are not homologous, they are analogue flight organs. However, the phalanges within them are homologous.

Chimpanzees have lower limbs that are specialized for manipulation, and (arguably) have fingers (instead of toes) on their lower limbs as well. In the case of primates in general, the digits of the hand are overwhelmingly referred to as "fingers". Primate fingers have both fingernails and fingerprints.

Research has been carried out on the embryonic development of domestic chickens showing that an interdigital webbing forms between the tissues that become the toes, which subsequently regresses by apoptosis. If apoptosis fails to occur, the interdigital skin remains intact. Many animals have developed webbed feet or skin between the fingers from this like the Wallace's flying frog.

Human fingers
Usually humans have five digits, the bones of which are termed phalanges, on each hand, although some people have more or fewer than five due to congenital disorders such as polydactyly or oligodactyly, or accidental or intentional amputations. The first digit is the thumb, followed by the index finger, middle finger, ring finger, and little finger or pinkie. According to different definitions, the thumb can be called a finger, or not.

English dictionaries describe finger as meaning either one of the five digits including the thumb, or one of the four digits excluding the thumb (in which case they are numbered from 1 to 4 starting with the index finger closest to the thumb).

Skeleton
The thumb (connected to the trapezium) is located on one of the sides, parallel to the arm.

The palm has five bones known as metacarpal bones, one to each of the five digits. Human hands contain fourteen digital bones, also called phalanges, or phalanx bones: two in the thumb (the thumb has no middle phalanx) and three in each of the four fingers. These are the distal phalanx, carrying the nail, the middle phalanx, and the proximal phalanx. Joints are formed wherever two or more of these bones meet. Each of the fingers has three joints:

 * metacarpophalangeal joint (MCP) – the joint at the base of the finger
 * proximal interphalangeal joint (PIP) – the joint in the middle of the finger
 * distal interphalangeal joint (DIP) – the joint closest to the fingertip.

Sesamoid bones are small ossified nodes embedded in the tendons to provide extra leverage and reduce pressure on the underlying tissue. Many exist around the palm at the bases of the digits; the exact number varies between different people.

The articulations are: interphalangeal articulations between phalangeal bones, and metacarpophalangeal joints connecting the phalanges to the metacarpal bones.

Muscles
Each finger may flex and extend, abduct and adduct, and so also circumduct. Flexion is by far the strongest movement. In humans, there are two large muscles that produce flexion of each finger, and additional muscles that augment the movement. The muscle bulks that move each finger may be partly blended, and the tendons may be attached to each other by a net of fibrous tissue, preventing completely free movement. Although each finger seems to move independently, moving one finger also moves the other fingers slightly which is called finger interdependence or finger enslaving.

Fingers do not contain muscles (other than arrector pili). The muscles that move the finger joints are in the palm and forearm. The long tendons that deliver motion from the forearm muscles may be observed to move under the skin at the wrist and on the back of the hand.

Muscles of the fingers can be subdivided into extrinsic and intrinsic muscles. The extrinsic muscles are the long flexors and extensors. They are called extrinsic because the muscle belly is located on the forearm.

The fingers have two long flexors, located on the underside of the forearm. They insert by tendons to the phalanges of the fingers. The deep flexor attaches to the distal phalanx, and the superficial flexor attaches to the middle phalanx. The flexors allow for the actual bending of the fingers. The thumb has one long flexor and a short flexor in the thenar muscle group. The human thumb also has other muscles in the thenar group (opponens and abductor brevis muscle), moving the thumb in opposition, making grasping possible.

The extensors are located on the back of the forearm and are connected in a more complex way than the flexors to the dorsum of the fingers. The tendons unite with the interosseous and lumbrical muscles to form the extensorhood mechanism. The primary function of the extensors is to straighten out the digits. The thumb has two extensors in the forearm; the tendons of these form the anatomical snuff box. Also, the index finger and the little finger have an extra extensor, used for instance for pointing. The extensors are situated within six separate compartments. The first compartment contains abductor pollicis longus and extensor pollicis brevis. The second compartment contains extensors carpi radialis longus and brevis. The third compartment contains extensor pollicis longus. The extensor digitorum indicis and extensor digitorum communis are within the fourth compartment. Extensor digiti minimi is in the fifth, and extensor carpi ulnaris is in the sixth.

The intrinsic muscle groups are the thenar and hypothenar muscles (thenar referring to the thumb, hypothenar to the small finger), the dorsal and palmar interossei muscles (between the metacarpal bones) and the lumbrical muscles. The lumbricals arise from the deep flexor (and are special because they have no bony origin) and insert on the dorsal extensor hood mechanism.

Skin
Aside from the genitals, the fingertips possess the highest concentration of touch receptors and thermoreceptors among all areas of the human skin, making them extremely sensitive to temperature, pressure, vibration, texture and moisture. A study in 2013 suggested fingers can feel nano-scale wrinkles on a seemingly smooth surface, a level of sensitivity not previously recorded. This makes the fingers commonly used sensory probes to ascertain properties of objects encountered in the world, making them prone to injury.

The  of a finger is the fleshy mass on the palmar aspect of the extremity of the finger.

Fingertip wrinkling in water
Although a common phenomenon, the underlying functions and mechanism of fingertip wrinkling following immersion in water are relatively unexplored. Originally it was assumed that the wrinkles were simply the result of the skin swelling in water, but it is now understood that the furrows are caused by the blood vessels constricting due to signalling by the sympathetic nervous system in response to water exposure. One hypothesis for why this occurs, the "rain tread" hypothesis, posits that the wrinkles may help the fingers grip things when wet, possibly being an adaption from a time when humans dealt with rain and dew in forested primate habitats. A 2013 study supporting this hypothesis found that the wrinkled fingertips provided better handling of wet objects but gave no advantage for handling dry objects. However, a 2014 study attempting to reproduce these results was unable to demonstrate any improvement of handling wet objects with wrinkled fingertips.

Regrowth of the fingertips
Fingertips, after having been torn off children, have been observed to regrow in less than 8 weeks. However, these fingertips do not look the same, although they do look more appealing than a skin graft or a sewn fingertip. No healing occurs if the tear happens below the nail. This works because the distal phalanges are regenerative in youth, and stem cells in the nails create new tissue that ends up as the fingertip.

Brain representation
Each finger has an orderly somatotopic representation on the cerebral cortex in the somatosensory cortex area 3b, part of area 1 and a distributed, overlapping representations in the supplementary motor area and primary motor area.

The somatosensory cortex representation of the hand is a dynamic reflection of the fingers on the external hand: in syndactyly people have a clubhand of webbed, shortened fingers. However, not only are the fingers of their hands fused, but the cortical maps of their individual fingers also form a club hand. The fingers can be surgically divided to make a more useful hand. Surgeons did this at the Institute of Reconstructive Plastic Surgery in New York to a 32-year-old man with the initials O. G.. They touched O. G.'s fingers before and after surgery while using MRI brain scans. Before the surgery, the fingers mapped onto his brain were fused close together; afterward, the maps of his individual fingers did indeed separate and take the layout corresponding to a normal hand.

Anomalies, injuries and diseases
A rare anatomical variation affects 1 in 500 humans, in which the individual has more than the usual number of digits; this is known as polydactyly. A human may also be born without one or more fingers or underdevelopment of some fingers such as symbrachydactyly. Extra fingers can be functional. One individual with seven fingers not only used them but claimed that they "gave him some advantages in playing the piano".

Phalanges are commonly fractured. A damaged tendon can cause significant loss of function in fine motor control, such as with a mallet finger. They can be damaged by cold, including frostbite and non-freezing cold injury (NFCI); and heat, including burns.

The fingers are commonly affected by diseases such as rheumatoid arthritis and gout. Individuals with diabetes often use the fingers to obtain blood samples for regular blood sugar testing. Raynaud's phenomenon and Paroxysmal hand hematoma are neurovascular disorders that affect the fingers.

Research has linked the ratio of lengths between the index and ring fingers to higher levels of testosterone, and to various physical and behavioral traits such as penis length and risk for development of alcohol dependence or video game addiction.

Etymology
The English word finger stems from Old English finger, ultimately from Proto-Germanic *fingraz ('finger'). It is cognate with Gothic figgrs, Old Norse fingr, or Old High German fingar. Linguists generally assume that *fingraz is a ro-stem deriving from a previous form *fimfe, ultimately from Proto-Indo-European *pénkʷe ('five').

The name pinkie derives from Dutch pinkje, of uncertain origin. In English only the digits on the hand are known as fingers. However, in some languages the translated version of fingers can mean either the digits on the hand or feet. In English a digit on a foot has the distinct name of toe.

title: Finger


text:
Talk:Anglo-Saxons/Archive 1

"collectively known as the Saxon conquest"? - In fact, it's more usually known as the Anglo-Saxon settlement(s); I've never heard "conquest" used before as a name/title (an event or process, yes, but always with a small "c"). Also, I don't think there's evidence of Roman abandonment before 407 (ie 5th C.), and more importantly, Angles weren't Saxons, they were Angles. User:David Parker My sentiments entirely. However, a number of historians refer to it as such, rightly or wrongly. It is very seldom referred to as the Anglo-Saxon settlement, and Anglo-Saxon bloodbath would be more altogether more apposite. Your settlement was anything but.

The Saxon Conquest is a slack generic name for the overlapping waves of Jutes, Danes, Angles, etc. that pitched up on our shores, and we can certainly counter this misnomer on these pages. It's provenance, however, is interesting: the Celtic population used the name "Saxon" generically to describe all the Germanic people they met. This indicates that there was a heavy preponderance of Saxons in the early raids and settlement. Please for one moment don't think the indigenous population gave up without a fight. The encroachment was not without its bloodshed at a level of savagery which might put the Israelis and the Palestinians to shame: Mons Badonicus, Dereham, Camulan, Otford Bridge, etc, etc, etc.

On, your second point, the Romans were never particularly interested in Britain, other than as a mechanism for suppressing the activities of the Druids and the Gauls, for whom Britain was a kind of early-day Afghanistan, a nice safe retreat from which to regroup, until the Romans took the country that is. The Roman empire's decline led to a progressive diminution of interest, and though it certainly accelerated duing the early 5th century, the process had started long before. There is plenty of documentary evidence for this. Certainly northern Britain was degarrisoned in about 300 AD after the troops were pulled back to help the empire fight back against barbarian incursions in central Europe, and Pictish incursions over the wall were many, frequent and bloody. sjc The term "Saxon" might have been used as a catch-all term among the invaders' opponents, but it was the Angles who gave their name to he northern kingdoms and eventually to the country as a whole; indeed "English settlement" is the other term used, which I'm again more familiar with than "Saxon Conquest".

The issue of a "bloodbath" is one that remains unresolved. The problem is that the only evidence for native population loss is our language, and there are other ways in which this could be accomplished, though the time-scale involved indeed suggests a less than gentle process. I imagine that then as later actual fighting involved only a small fraction of the population, most of whom had better things to do with their time, and I doubt that Anglo-Saxon military organisation made for very efficient executions of genocide, even if they'd been interested in such activities given the limited numbers which we can evisage wanting to migrate or being shifted across the sea. Most Britons I think probably indeed experienced settlement rather than warfare, though settlement "over" rather than "among" them.

I recognise that this view seems to be challenged by the recent BBC/UCL genetic survey, for which I've only seen very sketchy evidence so far, however. The whole convoluted German-Polish fracas illustrates how very different populations could co-exist, intermingle, exchange language and occasionally even embrace each other's rule to free themselves from some of their own "compatriots". Until mass British extinction is shown with some degree of certainty, I think the jury's still out on this one. User:David Parker

We need a sensible compromise name for this topic, and I don't think that I am going to sway you on this, and I have to say I have my own reservations.

But it must indicate that the settlement was definitely forced. Mons Badonicus was, even by modern standards, a major battle, and the splitting of the Cornish from the Welsh at the Battle of Deorham was a decisive moment, perhaps the decisive moment, in English history; had the tide of battle turned the other way, and by all accounts it could have done, this article would be being written in Brythonic Celtic.

The fighting was not only against the indigenous population; the internecine warfare between the settlers was on a massive scale: Offa conducted years of warfare against the Hestingii for example.

And yes, I have to admit the genetic survey was very interesting.

Where do you suggest removing this article to? User:sjc Hey, I'm swayable - I'm not out to start an argument, sjc, I just thought the issues were worth raising. As I said, the first genetic results look quite different from what I'd been expecting. I'm certainly not suggesting that settlement was pacific, and I don't think that's what others meant in the past. Personally, I'd recommend plain "Anglo-Saxons" as a title, which could start from discussion of their continental origins and conclude with modern uses of the term (de Gaulle et al). But I'm not out to interfere with your discussion of the conquest/settlement per se. Cheers, User:David Parker

I wasn't really suggesting you were. It's just an interesting little academic debate. Anglo-Saxons it is then. Cheers, sjc.

title: Talk:Anglo-Saxons/Archive 1


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User:Benwbrum

Ben W. Brumfield Austin, Texas

Occasional editor of Wikipedia since getting hooked in early 2002. Blogger and speaker about crowdsourcing and manuscript transcription. Inventor of the crowdsourced transcription tool FromThePage, which was largely inspired by Wikipedia. Best reached at benwbrum@gmail.com

See also User:Benwbrum/Hapiru for my comments on the Hapiru article.

Articles I've contributed to are (in decreasing order):
 * Tobacco
 * Akkadian language
 * Transliteration
 * Orange County, Texas
 * Texas Longhorn
 * Texas Revolution
 * Acadians
 * Semitic_languages
 * Cuneiform
 * Phonetic complement
 * Sorbian language
 * Elamites

title: User:Benwbrum