Holstege Gert, Subramanian Hari H
Asia-Pacific Centre for Neuromodulation, Queensland Brain Institute, The University of Queensland, Brisbane 4072, Australia.
J Comp Neurol. 2016 Jun 1;524(8):1558-77. doi: 10.1002/cne.23898. Epub 2015 Nov 17.
Vocalizations such as mews and cries in cats or crying and laughter in humans are examples of expression of emotions. These vocalizations are generated by the emotional motor system, in which the mesencephalic periaqueductal gray (PAG) plays a central role, as demonstrated by the fact that lesions in the PAG lead to complete mutism in cats, monkeys, as well as in humans. The PAG receives strong projections from higher limbic regions and from the anterior cingulate, insula, and orbitofrontal cortical areas. In turn, the PAG has strong access to the caudal medullary nucleus retroambiguus (NRA). The NRA is the only cell group that has direct access to the motoneurons involved in vocalization, i.e., the motoneuronal cell groups innervating soft palate, pharynx, and larynx as well as diaphragm, intercostal, abdominal, and pelvic floor muscles. Together they determine the intraabdominal, intrathoracic, and subglottic pressure, control of which is necessary for generating vocalization. Only humans can speak, because, via the lateral component of the volitional or somatic motor system, they are able to modulate vocalization into words and sentences. For this modulation they use their motor cortex, which, via its corticobulbar fibers, has direct access to the motoneurons innervating the muscles of face, mouth, tongue, larynx, and pharynx. In conclusion, humans generate speech by activating two motor systems. They generate vocalization by activating the prefrontal-PAG-NRA-motoneuronal pathway, and, at the same time, they modulate this vocalization into words and sentences by activating the corticobulbar fibers to the face, mouth, tongue, larynx, and pharynx motoneurons.
猫发出的喵叫声和哭声,或人类的哭泣和笑声等发声都是情感表达的例子。这些发声由情感运动系统产生,其中中脑导水管周围灰质(PAG)起着核心作用,这一事实表明,PAG损伤会导致猫、猴子以及人类完全缄默。PAG接受来自更高边缘区域以及前扣带回、岛叶和眶额叶皮质区域的强烈投射。反过来,PAG可以强烈影响延髓尾侧疑核(NRA)。NRA是唯一直接与参与发声的运动神经元相连的细胞群,即支配软腭、咽部、喉部以及膈肌、肋间肌、腹肌和盆底肌的运动神经元细胞群。它们共同决定腹内压、胸内压和声门下压力,控制这些压力对于发声是必要的。只有人类能够说话,因为通过意志性或躯体运动系统的外侧部分,他们能够将发声调节成单词和句子。为此,他们利用运动皮层,运动皮层通过其皮质延髓纤维直接与支配面部、口腔、舌头、喉部和咽部肌肉的运动神经元相连。总之,人类通过激活两个运动系统来产生言语。他们通过激活前额叶 - PAG - NRA - 运动神经元通路来产生发声,同时,他们通过激活通向面部、口腔、舌头、喉部和咽部运动神经元的皮质延髓纤维,将这种发声调节成单词和句子。