User:Psycho CSL/繮核

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韁核
韁核（紅色）位於松果體或丘腦前面的位置。
如圖，韁核（藍色）位於松果體（紅色）前面。
《神经解剖学术语（英语：Anatomical terms of neuroanatomy）》

韁核 （habenula，拉丁語中habena表示韁繩）是位於脊椎動物丘腦背側的一個體積較小的雙側核團，其體積小於一粒豌豆，形狀細長。其與第三腦室接壤，位於松果體前面。[1]

儘管韁核十分微小，但每個韁核都分為兩個不同的區域：內側韁核 (MHb) 和外側韁核 (LHb)，兩者都具有不同的神經元群、傳入纖維和傳出纖維。[2][3] 內側韁核可以再分為五個亞核，而外側韁核則可分為四個亞核。[4]研究顯示內側韁核及外側韁核形態的複雜性，內側韁核不同的傳入纖維分別投射到不同的亞核。[5]內外側韁核之間不同的基因表現使兩個區域有不同的功能。[6]

韁核是脊椎動物演化中的保守結構，哺乳動物的韁核是高度對稱的，而魚類、兩棲類及爬蟲類的韁核在大小、分子組成及連接方面都極不對稱。[1]韁核是邊緣系統通路中一個主要組成部分，[1]韁核和腳間核（英语：interpeduncular nucleus）之間的後屈束（英语：fasciculus retroflexus）路徑是發育中的大腦中首先形成的主要神經束之一。[1]

韁核是連接前腦區域和中腦區域的中央結構，為情緒和感覺處理整合的樞紐或交點，[2]它整合來自邊緣系統、感官和基底核的訊息來做出適當且有效的反應措施。[5]韁核參與單胺神經傳導物質的調節，特別是多巴胺和血清素，[2][3]這兩種神經傳導物質都與焦慮症和迴避行為密切相關。[2]韁核的功能也涉及動機、情緒、學習以及痛覺，[2]內側韁核在憂鬱、壓力、記憶和尼古丁戒斷中發揮重要作用，也在古柯鹼、安非他命和酒精成癮中發揮重要作用。[6]內側韁核表現出高水平的菸鹼型乙醯膽鹼受體 (nAChR)，其參與多種形式的成癮。[6]

解剖學[编辑]

每個韁核都分為內側及外側兩個部分，研究顯示內側韁核可以再分為五個亞核，而外側韁核則可分為四個亞核。[4] 左右韁核由韁連合（英语：habenular commissure）相連接，松果體附著於腦部此區域。[7]

外側韁核[编辑]

外側韁核主要的傳入區域是外側視前區（英语：preoptic area）、腹側蒼白球（英语：pallidum）、外側下視丘 The primary input regions to the lateral habenula (LHb) are the lateral preoptic area (bringing input from the hippocampus and lateral septum), the ventral pallidum (bringing input from the nucleus accumbens and mediodorsal nucleus of the thalamus), the lateral hypothalamus, the medial habenula, and the internal segment of the globus pallidus (bringing input from other basal ganglia structures).[8]

Neurons in the lateral habenula are 'reward-negative' as they are activated by stimuli associated with unpleasant events, the absence of the reward or the presence of punishment especially when this is unpredictable.[9] Reward information to the lateral habenula comes from the internal part of the globus pallidus.[10]

The outputs of the lateral habenula target dopaminergic regions (substantia nigra pars compacta and the ventral tegmental area), serotonergic regions (median raphe and dorsal raphe nuclei), and a cholinergic region (the laterodorsal tegmental nucleus).[8] This output inhibits dopamine neurons in substantia nigra pars compacta and the ventral tegmental area, with activation in the lateral habenula linking to deactivation in them, and vice versa, deactivation in the lateral habenula with their activation.[11] The lateral habenula functions to oppose the action of the laterodorsal tegmental nucleus in the acquisition of avoidance responses but not the processing of avoidance later on when it is a memory, motivation or its execution.[12] Research suggests that lateral habenula may play a crucial role in decision making.[13]

There has also shown to be an association with aberrant LHb activity and depression.[14]

內側韁核[编辑]

The medial habenula (MHb) receives connections from posterior septum pellucidum and diagonal band of Broca; the lateral habenula receives afferents from the lateral hypothalamus, nucleus accumbens, internal globus pallidus, ventral pallidum, and diagonal band of Broca.[8] As a whole, this complexly interconnected region is part of the dorsal diencephalic conduction system (DDCS), responsible for relaying information from the limbic system to the midbrain, hindbrain, and medial forebrain.[15][16]

Input to the medial habenula comes from a variety of regions and carries a number of different chemicals. Input regions include septal nuclei (the nucleus fimbrialis septi and the nucleus triangularis septi), dopaminergic inputs from the interfascicular nucleus of the ventral tegmental area, noradrenergic inputs from the locus ceruleus, and GABAergic inputs from the diagonal band of Broca. The medial habenula sends outputs of glutamate, substance P and acetylcholine to the periaqueductal gray via the interpeduncular nucleus as well as to the pineal gland.[17][18]

Asymmetry[编辑]

Nikolaus Goronowitsch[7]在1883年發現了韁核的不對稱性，許多物種皆展現了韁核神經元左右不對稱的分化。 Asymmetry in the habenula was first noted in 1883 by Nikolaus Goronowitsch.[7] Various species exhibit left-right asymmetric differentiation of habenular neurons. In many fishes and amphibians, the habenula on one side is significantly larger and better organized into distinct nuclei in the dorsal diencephalon than its smaller pair. The sidedness of such differentiation (whether the left or the right is more developed) varies with the species. In birds and mammals, however, both habenulae are more symmetrical (although not entirely) and consist of a medial and a lateral nucleus on each side which is in fish and amphibians equivalent to dorsal habenula and the ventral habenula, respectively.[19][8][20]

Olfactory coding[编辑]

In some fish (lampreys and teleosts), mitral cell (principal olfactory neurons) axons project exclusively to the right hemisphere of the habenula in an asymmetric manner. It is reported that the dorsal habenula (DHb) are functionally asymmetric with predominantly odor responses in the right hemisphere. It was also shown that DHb neurons are spontaneously active even in the absence of olfactory stimulation. These spontaneously-active DHb neurons are organized into functional clusters which were proposed to govern olfactory responses. (Jetti, SK. et al 2014, Current Biology)

功能[编辑]

These nuclei are hypothesized to be involved in regulation of monoamines, such as dopamine and serotonin.[21][22]

The habenular nuclei are involved in pain processing, reproductive behavior, nutrition, sleep-wake cycles, stress responses, and learning. Recent demonstrations using fMRI[23] and single unit electrophysiology[11] have closely linked the function of the lateral habenula with reward processing, in particular with regard to encoding negative feedback or negative rewards. Matsumoto and Hikosaka suggested in 2007 that this reward and reward-negative information in the brain might "be elaborated through the interplay among the lateral habenula, the basal ganglia, and monoaminergic (dopaminergic and serotonergic) systems" and that the lateral habenula may play a pivotal role in this "integrative function".[11] Then, Bromberg-Martin et al. (2011) highlighted that neurons in the lateral habenula signal positive and negative information-prediction errors in addition to positive and negative reward-prediction errors.[24]

與憂鬱症的關聯[编辑]

Both the medial and lateral habenula show reduced volume in those with depression. Neuron cell numbers were also reduced on the right side.[25] Such changes are not seen in those with schizophrenia.[25] Deep brain stimulation of the major afferent bundle (i.e., stria medullaris thalami) of the lateral habenula has been used for treatment of depression where it is severe, protracted and therapy-resistant.[26][27]

N-Methyl-D-aspartate (NMDA) receptor-dependent burst firing in the lateral habenula has been associated with depression in animal studies,[28] and it has been shown that the general anesthetic ketamine blocks this firing by acting as a receptor antagonist.[29] Ketamine has been the subject of numerous studies after having shown fast-acting antidepressant effects in humans (in a 0.5 mg/bw kg dose).[30]

Motivation and addiction[编辑]

Recent exploration of the habenular nuclei has begun to associate the structure with an organism's current mood, feeling of motivation, and reward recognition.[31] Previously, the LHb has been identified as an "anti-reward" signal, but recent research suggests that the LHb helps identify preference, helping the brain to discriminate between potential actions and subsequent motivation decisions.[32] In a study using a Pavlovian conditioning model, results showed an increase in the habenula response.[33] This increase coincided with conditioned stimuli associated with more aversive punishments (i.e. electric shock).[33] Therefore, researchers speculate that inhibition or damage to the LHb resulting in a failure to process such information may lead to random motivation behavior.[32][33]LHb is especially important in understanding the reward and motivation relationship as it relates to addictive behaviors.[31] The LHb inhibits dopaminergic neurons, decreasing the release of dopamine.[34] It was determined by several animal studies that receiving a reward coincided with elevated dopamine levels, but once the learned association was learned by the animal, dopamine levels remain elevated, only decreasing when the reward is removed.[20][22][31][34] Therefore, dopamine levels only increase with unpredicted rewards and with a "positive prediction error".[20] Moreover, it was determined that removal of an anticipated award activated LHb, inhibited dopamine levels.[20] This finding helps explain why addictive drugs are associated with elevated dopamine levels.[20]

Nicotine and nAChRs[编辑]

According to the National Institute on Drug Abuse, one in five preventable deaths in the United States is caused by tobacco use.[35] Nicotine is the addictive drug found in most tobacco products and is easily absorbed by the bloodstream of the body.[35] Despite common misconceptions regarding the relaxing effects of tobacco and nicotine use, behavioral testing in animals has demonstrated nicotine to have an anxiogenic effect.[36] Nicotinic acetylcholine receptors (nAChRs) have been identified as the primary site for nicotine activity and regulate consequent cellular polarization.[37] nAChRs are made up a number of α and β subunits and are found in both the LHb and MHb, where research suggests they may play a key role in addiction and withdrawal behaviors.[37][38]

歷史[编辑]

韁核是一種在3.6億多年前出現在脊椎動物中的保守結構，[4]安德烈亞斯·維薩留斯於1555年首次描述了韁連合，[39]而狄奧多·赫曼·梅涅特則在1872年提到了韁核。[40]

參考來源[编辑]

  1. ^ 1.0 1.1 1.2 1.3 Antolin-Fontes, B; Ables, JL; Görlich, A; Ibañez-Tallon, I. The habenulo-interpeduncular pathway in nicotine aversion and withdrawal.. Neuropharmacology. September 2015, 96 (Pt B): 213–22. PMC 4452453 可免费查阅. PMID 25476971. doi:10.1016/j.neuropharm.2014.11.019. 
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  3. ^ 3.0 3.1 Boulos, LJ; Darcq, E; Kieffer, BL. Translating the Habenula-From Rodents to Humans.. Biological Psychiatry. 15 February 2017, 81 (4): 296–305. PMC 5143215 可免费查阅. PMID 27527822. doi:10.1016/j.biopsych.2016.06.003. 
  4. ^ 4.0 4.1 4.2 Ables, JL; Park, K; Ibañez-Tallon, I. Understanding the habenula: A major node in circuits regulating emotion and motivation.. Pharmacological Research. April 2023, 190: 106734. PMID 36933754. doi:10.1016/j.phrs.2023.106734 可免费查阅. 
  5. ^ 5.0 5.1 Juárez-Leal, I; Carretero-Rodríguez, E; Almagro-García, F; Martínez, S; Echevarría, D; Puelles, E. Stria medullaris innervation follows the transcriptomic division of the habenula.. Scientific Reports. 16 June 2022, 12 (1): 10118. Bibcode:2022NatSR..1210118J. PMC 9203815 可免费查阅. PMID 35710872. doi:10.1038/s41598-022-14328-1. 
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  18. ^ Antolin-Fontes B, Ables JL, Görlich A, Ibañez-Tallon I. The habenulo-interpeduncular pathway in nicotine aversion and withdrawal. Neuropharmacology. September 2015, 96 (Pt B): 213–22. PMC 4452453 可免费查阅. PMID 25476971. doi:10.1016/j.neuropharm.2014.11.019. 
  19. ^ Hüsken U, Stickney HL, Gestri G, Bianco IH, Faro A, Young RM, Roussigne M, Hawkins TA, Beretta CA, Brinkmann I, Paolini A, Jacinto R, Albadri S, Dreosti E, Tsalavouta M, Schwarz Q, Cavodeassi F, Barth AK, Wen L, Zhang B, Blader P, Yaksi E, Poggi L, Zigman M, Lin S, Wilson SW, Carl M. Tcf7l2 is required for left-right asymmetric differentiation of habenular neurons. Current Biology. October 2014, 24 (19): 2217–27. PMC 4194317 可免费查阅. PMID 25201686. doi:10.1016/j.cub.2014.08.006. 
  20. ^ 20.0 20.1 20.2 20.3 20.4 Hu, Hailan; Cui, Yihui; Yang, Yan. Circuits and functions of the lateral habenula in health and in disease. Nature Reviews Neuroscience. April 2020, 21 (5): 277–295. ISSN 1471-0048. PMID 32269316. S2CID 215411587. doi:10.1038/s41583-020-0292-4 （英语）. 
  21. ^ Stephenson-Jones, Marcus; Floros, Orestis; Robertson, Brita; Grillner, Sten. Evolutionary conservation of the habenular nuclei and their circuitry controlling the dopamine and 5-hydroxytryptophan (5-HT) systems. Proceedings of the National Academy of Sciences of the United States of America. 2011, 109 (3): E164–E173. PMC 3271889 可免费查阅. PMID 22203996. doi:10.1073/pnas.1119348109 可免费查阅. 
  22. ^ 22.0 22.1 Boulos, Laura-Joy; Darcq, Emmanuel; Kieffer, Brigitte Lina. Translating the Habenula—From Rodents to Humans. Biological Psychiatry. 2017-02-15, 81 (4): 296–305. PMC 5143215 可免费查阅. PMID 27527822. doi:10.1016/j.biopsych.2016.06.003. 
  23. ^ Ullsperger M, von Cramon DY. Error monitoring using external feedback: specific roles of the habenular complex, the reward system, and the cingulate motor area revealed by functional magnetic resonance imaging. The Journal of Neuroscience. May 2003, 23 (10): 4308–14. PMC 6741115 可免费查阅. PMID 12764119. doi:10.1523/JNEUROSCI.23-10-04308.2003. 
  24. ^ Bromberg-Martin ES, Hikosaka O. Lateral habenula neurons signal errors in the prediction of reward information. Nature Neuroscience. August 2011, 14 (9): 1209–16. PMC 3164948 可免费查阅. PMID 21857659. doi:10.1038/nn.2902. 
  25. ^ 25.0 25.1 Ranft K, Dobrowolny H, Krell D, Bielau H, Bogerts B, Bernstein HG. Evidence for structural abnormalities of the human habenular complex in affective disorders but not in schizophrenia. Psychological Medicine. April 2010, 40 (4): 557–67. PMID 19671211. S2CID 11799795. doi:10.1017/S0033291709990821. 
  26. ^ Sartorius A, Kiening KL, Kirsch P, von Gall CC, Haberkorn U, Unterberg AW, Henn FA, Meyer-Lindenberg A. Remission of major depression under deep brain stimulation of the lateral habenula in a therapy-refractory patient. Biological Psychiatry. January 2010, 67 (2): e9–e11. PMID 19846068. S2CID 43590983. doi:10.1016/j.biopsych.2009.08.027. 
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  33. ^ 33.0 33.1 33.2 Lawson, Rebecca P.; Seymour, Ben; Loh, Eleanor; Lutti, Antoine; Dolan, Raymond J.; Dayan, Peter; Weiskopf, Nikolaus; Roiser, Jonathan P. The habenula encodes negative motivational value associated with primary punishment in humans. Proceedings of the National Academy of Sciences. 2014-08-12, 111 (32): 11858–11863. Bibcode:2014PNAS..11111858L. ISSN 0027-8424. PMC 4136587 可免费查阅. PMID 25071182. doi:10.1073/pnas.1323586111 可免费查阅 （英语）. 
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外部連結[编辑]


  • Jetti SK, Vendrell-Llopis N, Yaksi E. Spontaneous activity governs olfactory representations in spatially organized habenular microcircuits. Current Biology. February 2014, 24 (4): 434–9. PMID 24508164. doi:10.1016/j.cub.2014.01.015 可免费查阅. 


人類间脑解剖
上丘脑
表面
灰质
丘脑
表面
灰质/
nuclei（英语：List of thalamic nuclei）
白質
下丘脑
表面
灰质
自主神经系统
內分泌系統
情绪
白質
垂體
底丘脑
取自“https://zh.wikipedia.org/w/index.php?title=User:Psycho_CSL/繮核&oldid=82118479”