Neurophysiology

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Neurophysiology is a branch of physiology and neuroscience concerned with the functions of the nervous system and their mechanisms. The term neurophysiology originates from the Greek word νεῦρον ("nerve") and physiology (which is, in turn, derived from the Greek φύσις, meaning "nature", and -λογία, meaning "knowledge").<ref name = "Luhmann 2013"></ref> Neurophysiology has applications in the prevention, diagnosis, and treatment of many neurological and psychiatric diseases.<ref name = "Delvendahl & Hu 2023"/> Neurophysiological techniques are also used by clinical neurophysiologists to diagnose and monitor patients with neurological diseases.

The field involves all levels of nervous system function, from molecules and cells to systems and whole organisms. Areas of study include:

Experimental neurophysiologists use many techniques to study neural function. Electrophysiological techniques like electroencephalography (EEG), single cell recording, and extracellular recording of local field potentials are especially common.<ref name = "Phillipu 2017/> Multi-electrode arrays on semiconductor chips can perform in vitro extracellular recording<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> and in vitro intracellular recording<ref>Template:Cite journal</ref> at scale. Magnetoencephalography is sometimes used in place of EEG.<ref name = "Stam & Straaten 2012"/> Immunohistochemistry, cell staining, in situ hybridisation, calcium imaging, and transmission electron microscopy are used to study cellular activity in the nervous system. Genetic engineering techniques may be used to study the impact of specific genes on neural functions. Pharmacological methods are used investigate the function of specific receptors in neurons and glia. Optogenetics and chemogenetics allow specific activation of neurons to study their functions. Functional magnetic resonance imaging and positron emission tomography can be used to measure metabolic changes in the brain.<ref name = "Giovacchini et al 2010" /><ref name = "Stam & Straaten 2012"/> Finally, behavioural analysis is used to understand interactions between physiology and behaviour. Contemporary neurophysiology experiments often use multiple techniques together to develop a more complete understanding of their research areas.

History

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Antiquity

File:Images from Edwin Smith Papyrus Wellcome L0003140.jpg
The Breasted edition (1930): left page photograph of the original Edwin Smith papyrus, right page transcription of hieroglyphics. This is Plate XIII (column 13, case 38-41)

Neurophysiology has been a subject of study since as early as 4,000 B.C. In the early B.C. years, most studies were of different natural sedatives like alcohol and poppy plants.Template:Citation needed In 1700 B.C., the Edwin Smith surgical papyrus was written. This papyrus was crucial in understanding how the ancient Egyptians understood the nervous system. This papyrus looked at different case studies about injuries to different parts of the body, most notably the head. The knowledge presents a rational and scientific approach to medicine in ancient Egypt.<ref name="Ritner 2001">Ritner, R.K. (2001). "Magic. In The Oxford Encyclopedia of Ancient Egypt". : Oxford University Press. Retrieved 8 Nov. 2025, from https://www.oxfordreference.com/view/10.1093/acref/9780195102345.001.0001/acref-9780195102345-e-0424.</ref>Template:Rp Beginning around 460 B.C., Hippocrates began to study epilepsy, and theorized that it had its origins in the brain. Hippocrates also theorized that the brain was involved in sensation, and that it was where intelligence was derived from. Hippocrates, as well as most ancient Greeks, believed that relaxation and a stress free environment was crucial in helping treat neurological disorders, that "healthy Mediterranean diet and daily moderate physical activity can prevent disease".<ref name="Hippocrates_Kostakopoulos">Template:Cite journal</ref> In 280 B.C., Erasistratus of Chios theorized that there were divisions in vestibular processing in the brain, as well as deducing from observation that sensation was located there.<ref name="Wills et al 1999">Wills, Adrian, and A Wills. “Herophilus, Erasistratus, and the Birth of Neuroscience.” Lancet 354, no. 9191 (November 13, 1999): 1719–20. doi:10.1016/S0140-6736(99)02081-4.</ref>

In 177 Galen theorized that human thought occurred in the brain, as opposed to the heart as Aristotle had theorized.<ref>Template:Cite journal</ref> The optic chiasm, which is crucial to the visual system, was discovered around 100 C.E. by Marinus.

Middle ages

Abū Bakr al-Rāzī (864 or 865–925 or 935 C.E.), also known by his Latin name Rhazes, a Persian physician, philosopher, and alchemist, wrote a book for al-Mansur (Kitāb al-Manṣūrī) of 26 sections on body structures, including nerves, muscles, and organs such as the eyes and heart.<ref>Template:Cite encyclopedia</ref><ref>Walker, Paul E. (1998). "al-Razi, Abu Bakr Muhammad ibn Zakariyya' (d. 925)". In Craig, Edward (ed.). Routledge Encyclopedia of Philosophy. doi:10.4324/9780415249126-H043-1. ISBN 978-0-415-25069-6.</ref><ref>Alghamdi MA, Ziermann JM, Diogo R. (2017). "An untold story: The important contributions of Muslim scholars for the understanding of human anatomy". The Anatomical Record. 2017 Jun;300(6):986-1008.</ref> It was translated into Latin by Gerard of Cremona around 1180.<ref>Template:Cite web</ref> Template:Circa, Al-Zahrawi, living in Iberia, began to write about different surgical treatments for neurological disorders. He described a surgical procedure for treating such a neurological disease as migraine (in particular, ligation of the temporal artery for migraine),<ref>Template:Cite journal</ref> in his thirty-volume medical encyclopedia, the Kitab al-Tasrif manuscript, completed in the year 1000.<ref>Template:Cite journal</ref> Al-Zahrawi was first to discover the root cause of paralysis.<ref name="MedievalMedicine">Template:Cite book</ref> In Persia, Avicenna (Ibn-Sina) presented detailed knowledge about skull fractures and their surgical treatments.<ref>Template:Cite journal</ref> In 1216, the first anatomy textbook in Europe, which included a description of the brain, was written by Mondino de Luzzi.<ref>Wilson, Luke (1987). "The performance of the body in the Renaissance theater of anatomy". p.64. Representations (17): 62–95.</ref>Template:Rp In 1402, St Mary of Bethlehem Hospital (later known as Bedlam in Britain) was the first hospital used exclusively for the mentally ill.<ref>Template:Cite web</ref>

16th century

In 1504, Leonardo da Vinci continued his study of the human body with a wax cast of the human ventricle system. In 1536, Nicolo Massa described the effects of different diseases, such as syphilis on the nervous system. He also noticed that the ventricular cavities were filled with cerebrospinal fluid.<ref>Template:Cite book</ref> In 1542, the term physiology was used for the first time by a French physician named Jean Fernel, to explain bodily function in relation to the brain.<ref>Tubbs, S.R. (2015). "Anatomy is to physiology as geography is to history; it describes the theatre of events". Clinical Anatomy. 28 (2): 151. https://doi.org/10.1002/ca.22526</ref>

File:Vesalius 609c.png
Vesalius arranged his work into seven books. Vesalius' De humani corporis fabrica, Book 7: The Brain, figure on plate 609, contrasted.

In 1543, Andreas Vesalius wrote De humani corporis fabrica, which revolutionized the study of anatomy. The book 7 of this corpus described the structure and functions of the brain and its coverings, the eye, the organs of sensation, and the nerves of the limbs. In this book, he described the pineal gland and what he believed the function was, and was able to draw the corpus striatum which is made up of the basal ganglia and the internal capsule. It was closed with a chapter on the correct way to dissect the brain.<ref>Template:Cite journal</ref> In 1549, Jason Pratensis published De Cerebri Morbis. This book was devoted to neurological diseases, and discussed symptoms, as well as ideas from Galen and other Greek, Roman and Arabic authors. It also looked into the anatomy and specific functions of different areas.<ref>Template:Cite journal</ref> In 1550, Andreas Vesalius worked on a case of hydrocephalus, or fluid filling the brain. In the same year, Bartolomeo Eustachi studied the optic nerve, mainly focusing on its origin in the brain. In 1564, Giulio Cesare Aranzio discovered the hippocampus,<ref>Gurunluoglu, R; Shafighi, M.; Gurunluoglu, A. (2011) "Giulio Cesare Aranzio (Arantius) (1530–89) in the pageant of anatomy and surgery". Journal of Medical Biography, vol. 19, n. 2, 2011-05, pp. 63-69, https://doi.org/10.1258/jmb.2010.010049</ref> naming it such due to its shape resemblance to a sea horse.

17th century

In 1621, Robert Burton published The Anatomy of Melancholy, which looked at the loss of important characters in one's life as leading to depression.<ref>Template:Cite book</ref> In 1649, René Descartes studied the pineal gland. He mistakenly believed that it was the "soul" of the brain, and believed it was where thoughts formed. In 1658, Johann Jakob Wepfer studied a patient in which he believed that a broken blood vessel had caused apoplexy, or a stroke. The European Stroke Conference has awarded the Wepfer Prize for stroke research annually since 2005.<ref>Template:Cite journal</ref> In 1664, Thomas Willis published his Anatomy of the Brain. He described the brain more clearly, setting forth the circle of Willis, the circle of vessels that enables arterial supply of the brain. He described epilepsy, apoplexy, and paralysis.<ref>Template:Cite journal</ref>

18th century

In 1749, David Hartley published Observations on Man, which focused on frame (neurology), duty (moral psychology) and expectations (spirituality) and how these integrated within one another.<ref>Harry Whitaker, C. U. M.; Smith, Stanley Finger (eds.), (2007). Brain, Mind and Medicine: Essays in Eighteenth-Century Neuroscience, Springer, 2007, p. 177.</ref> This text was also the first to use the English term psychology. In 1752, the Society of Friends created an asylum in Philadelphia, Pennsylvania. The asylum intended to give not only medical treatment to those mentally ill, but also provide with caretakers and comfortable living conditions.Template:Citation needed In 1755, Jean-Baptiste Le Roy began using electroconvulsive therapy for the mentally ill, a treatment still used today in specific cases. In 1760, Arne-Charles studied how different lesions in the cerebellum could affect motor movements.<ref>Finger, S.(1994). Origins of Neuroscience. A History of Explorations into Brain Function. New York: Oxford University Press.</ref> In 1776, Template:Interlanguage link multi studied the cerebellum intensely, and published a book solely based on its function and appearance.

In 1784, Félix Vicq-d'Azyr, discovered a black colored structure in the midbrain.<ref name=" Tubbs11">Template:Cite journal</ref> In 1791 Samuel Thomas von Sömmerring alluded to this structure, calling it the substantia nigra.<ref>Swanson, LW. Neuroanatomical terminology : a lexicon of classical origins and historical foundations. Oxford University Press, 2014. England Template:ISBN</ref> In the same year, Luigi Galvani described the role of electricity in nerves of dissected frogs.<ref>Template:Cite book</ref>

19th century

In 1808, Franz Joseph Gall studied and published work on phrenology. Phrenology was the faulty science of looking at head shape to determine different aspects of personality and brain function.<ref>Bear, Mark F.; Connors, Barry W.; Paradiso, Michael A. (2007). Neuroscience. Lippincott Williams & Wilkins. ISBN 978-0-7817-6003-4.</ref>Template:Rp In 1811, Julien Jean César Legallois studied respiration in animal dissection and lesions and found the center of respiration in the medulla oblongata.<ref>Cheung, T. (2013). "Limits of Life and Death: Legallois’s Decapitation Experiments". J Hist Biol 46, 283–313 (2013). https://doi.org/10.1007/s10739-012-9335-7</ref> In the same year, Charles Bell finished work on what would later become known as the Bell–Magendie law, which compared functional differences between dorsal and ventral roots of the spinal cord. He was the first to distinguish between motor and sensory nerves and proposed the concept of a “muscle sense”.<ref>Conti, A.A. (2014). "Western medical rehabilitation through time: a historical and epistemological review". TheScientificWorldJournal, 2014, 432506. https://doi.org/10.1155/2014/432506</ref> In 1822, Karl Friedrich Burdach distinguished between the lateral and medial geniculate bodies, as well as named the cingulate gyrus. The column of Burdach or fasciculus cuneatus, the lateral portion of the dorsal funiculus of the spinal cord is named for him.<ref>The American Illustrated Medical Dictionary, 1938.</ref> In 1824, F. Magendie studied and produced the first evidence of the cerebellum's role in equilibration to complete the Bell–Magendie law. In 1838, Theodor Schwann began studying white and grey matter in the brain, and discovered the myelin sheath. These cells, which cover the axons of the neurons in the brain, are named Schwann cells after him.<ref name="Axel Karenberg">Template:Cite book</ref> In 1843 Carlo Matteucci and Emil du Bois-Reymond demonstrated that nerves transmit signals electrically.<ref>Template:Cite web</ref>

A diagram of Gage's skull
The Phineas Gage case: The left frontal lobe (red), with Ratiu et al.'s estimate of the iron's path

In 1848, Phineas Gage, the classical neurophysiology patient, had his brain pierced by an iron tamping rod in a blasting accident. He became an excellent case study in the connection between the prefrontal cortex and behavior, decision making and consequences.<ref>Bigelow, Henry Jacob (July 1850). "Dr. Harlow's Case of Recovery from the Passage of an Iron Bar through the Head". American Journal of the Medical Sciences. New series. 20 (39): 13–22.</ref>Template:Rp In 1849, Hermann von Helmholtz studied the speed of frog nerve impulses while studying electricity in the body.<ref name="Ian Glynn">Template:Cite book</ref>

In 1861, French neurologist Paul Broca discovered that a damaged area of the posterior inferior frontal gyrus also known as Broca's area) in patients caused an inability to speak.<ref>Template:Cite journal</ref>

Italian neuroanatomist professor Camillo Golgi discovered in the 1870s that all the nerve cells in the nervous system are a continuous, interconnected network.<ref>"Camillo Golgi – Facts". NobelPrize.org. Nobel Prize Outreach AB 2025. Tue. 14 Jan 2025. https://www.nobelprize.org/prizes/medicine/1906/golgi/facts/</ref>

Regionally specific features of the central brain region were described by Prof. Betz in 1874, making one of the first steps toward the microstructural, cytoarchitectonic parcellation of the entire human cerebral cortex.<ref>Solodkin, A.; Hlustik, P.; Buccino, G. (2007). "22 The Anatomy and Physiology of the Motor System" Retrieved 2025.11.15. chrome-extension://efaidnbmnnnibpcajpcglclefindmkaj/http://fmri.upol.cz/webdoc/solodkin-hlustik-buccino-motorChapterProof06.pdf</ref><ref>Betz, W. (1874). "Anatomischer Nachweis zweier Gehirncentra". Centralblatt fur die medizinische Wissenschaften, 12, 578–580, 595–599.</ref>

In 1875, Prof. Richard Caton reported to the British Medical Association that he had observed electrical impulses from the surfaces of living brains in animals.<ref>Finger, Stanley (1994). Origins of Neuroscience: a history of explorations in brain function. New York: Oxford University Press, pp. 41–42</ref>

In 1894, neurologist and psychiatrist Edward Flatau published a human brain atlas “Atlas of the Human Brain and the Course of the Nerve-Fibres”.<ref>Template:Cite journal</ref>

In 1896, Prof. d'Arsonval conducted the first documented study of the effects of a time-variable magnetic field on the brain, producing physiological changes: a volunteer reported phosphenes and vertigo when a coil stimulated his head at 42 Hz.<ref>Kobayashi, M.; Pascual-Leone, A.(2003). Transcranial magnetic stimulation in neurology. Lancet Neurol, 2(3), 145–156.</ref>

20th century

In 1902, Prof. Julius Bernstein contributed to the physiology of nerve impulses by proposing that the action potential results from a change in the permeability of the axonal membrane to ions, thereby providing knowledge of the origin of the "resting potential" and the "action potential" in the nerve. The "membrane hypothesis" explained the resting potential of nerve and muscle as a diffusion potential set up by the tendency of positively charged ions to diffuse from their high concentration in cytoplasm to their low concentration in the extracellular solution, while other ions are held back. Bernstein was also the first to introduce the Nernst equation for resting potential across the membrane.<ref>Seyfarth E-A. (2006), "Julius Bernstein (1839–1917): pioneer neurobiologist and biophysicist" Template:Webarchive, Biological Cybernetics 94: 2–8 Biol Cybern (2006) 94: 2–8 Template:Doi</ref>

In 1907, Prof. Louis Lapicque proposed that the action potential occurs at threshold crossing[39] (later better shown to be a product of dynamic ionic conduction systems).<ref>Template:Cite journal</ref>

File:Vergleichende Lokalisationslehre der Grosshirnrinde in ihren Prinzipien dargestellt auf Grund des Zellenbaues.pdf
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In 1909, German anatomist Korbinian Brodmann published his original research on brain mapping, defining 52 distinct regions of the cerebral cortex, known as Brodmann areas now.<ref>Template:Cite journal</ref>

In 1924, German physiologist and psychiatrist Hans Berger (1873–1941) discovered the electrical activity of the brain (called brain waves) and, in particular, the alpha wave rhythm, which is a type of brain wave.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

A great deal of study on sensory organs and the function of nerve cells was conducted by British physiologist Edgar Adrian. In 1928, he observed nerve fibers in action during his experiments on frogs, and was able to record the electrical discharge of single nerve fibres under physical stimulus.<ref name="Adrian">Template:Cite web</ref> He concluded that the excitation of the skin under constant stimulus is initially strong but gradually decreases over time, whereas the sensory impulses passing along the nerves from the point of contact are constant in strength, yet are reduced in frequency over time, and the sensation in the brain diminishes as a result.<ref name="Adrian"/> Extending these results to the study of pain causes by the stimulus of the nervous system, he made discoveries about the reception of such signals in the brain and spatial distribution of the sensory areas of the cerebral cortex in different animals. These conclusions lead to the idea of a sensory map, called the homunculus, in the somatosensory system. This distorted view of the human body was based on the neurological "map" of the regions and parts of the human brain response.<ref name="Adrian"/>

In 1944, Prof. Josef Erlanger and Herbert Gasser were awarded the Nobel Prize in Physiology or Medicine "for their discoveries relating to the highly differentiated functions of single nerve fibres."<ref>Template:Cite web</ref> They discovered action potentials of two phases: spike and postspike; and they revealed nerves in many forms, each with its own excitability potential. They also found a proportion between nerve fiber diameter and action potential velocity. <ref> Grant, Gunnar (2006). "The 1932 and 1944 Nobel Prizes in physiology or medicine: rewards for ground-breaking studies in neurophysiology". Journal of the History of the Neurosciences. 15 (4): 341–357. doi:10.1080/09647040600638981 </ref>

In 1950, Prof. Wilder Penfield published a book called The Cerebral Cortex of Man with maps of the location of various functions (motor, sensory, memory, vision) in the brain, that he defined during the process of treating epilepsy.<ref> Kumar, R.; Yeragani, V. K. (2011). "Penfield – A great explorer of psyche-soma-neuroscience". Indian Journal of Psychiatry. 53 (3): 276–278. doi:10.4103/0019-5545.86826 </ref> Prof. Wilder Penfield and his colleagues Edwin Baldry and Theodore Rasmussen are considered the creators of the cortical homunculus.<ref> Cazala, Fadwa; Vienney, Nicolas; Stoléru, Serge (2015-03-10). "The cortical sensory representation of genitalia in women and men: a systematic review". Socioaffective Neuroscience & Psychology. 5 26428. doi:10.3402/snp.v5.26428 </ref>

See also

References

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Sources

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