摘要
为了探讨牦牛脑神经对青藏高原生态环境适应的形态学机制,对青藏高原牦牛脑神经的起源、分布、分支等进行了全面、细致地解剖学研究。为进一步在分子、基因水平上探讨青藏高原牦牛的适应性机制及研究和开发青藏高原牦牛的优势遗传资源提供基础资料,最终加速优化青藏高原高寒草地生态系统的服务功能一生产功能的实现、为人类神经系统及高原病的研究和防治提供理论依据、促进我国西部地区社会经济和生态的可持续发展、填补国内外特有家畜研究领域的空白,特进行此项研究。主要的研究结果如下:
1.嗅神经起于鼻中隔后部以及上鼻道后部的粘膜,终止于嗅球的前腹侧面;终神经起于上鼻道后上部粘膜及鼻中隔后上部嗅区上皮的血管和腺体,终止于嗅三角内侧面和压胼胝体区域;犁鼻神经起于鼻腔前部底壁内侧的犁鼻器及其相邻区域及鼻中隔的粘膜,终止于嗅三角前内侧缘的嗅回处。
2.视神经起于眼球的视网膜视神经盘,穿越视神经管进入颅腔,在颅腔内斜向内侧延伸约2～3 cm后,终止于鞍隔背侧面的视交叉。视神经呈圆柱状,直径为4.60～5.50 mm,自视交叉至眼球视神经直径逐渐变细且相差约0.5mm。视神经在眼眶内呈“S”形延伸,且延伸途中汇入了1～2条睫状短神经。
3.动眼神经起于中脑的大脑脚腹面之后部脚间窝,经眶圆孔入眼眶后分为背侧支和腹侧支,主要分布于除眼外侧和背侧斜肌之外的所有眼球肌和上睑提肌,而且腹侧支还分出一条睫状神经汇入睫状神经节。
4.滑车神经起于四叠体后丘和前髓帆之间的沟内,经眶圆孔入眼眶,分布于眼背侧斜肌,并有交通支与眼神经相连。
5.三叉神经起于脑桥的背外侧,分为眼神经、上颌神经和下颌神经。
眼神经的分支有颧颞支、泪脉神经、额神经、额窦神经、肌支、鼻睫神经以及连于鼻睫神经节的交通支。鼻睫神经分为滑车下神经和筛神经。筛神经分为鼻内支和鼻外支。鼻睫神经有交通支连于睫状神经节。
上颌神经分出颧面支、副颧面支、颧骨神经、鼻后神经、腭大神经、腭小神经和上齿槽后支,其主干延续为眶下神经。上颌神经有较粗的交通支与眼神经和颊神经相连。鼻后神经在冀腭窝内分为数支参于构成翼腭神经丛。腭大神经分出腭副神经以及进入鼻腔底壁的侧支。
下颌神经分出颞深神经、咬肌神经、颊神经、舌神经、下齿槽神经、下颌舌骨肌神经、翼内侧肌神经、翼外侧肌神经和耳颞神经。颊神经有交通支与上颌神经、腭大神经、面神经颊背侧支相连,且有分支分布于翼内侧肌。下齿槽神经在下齿槽管内分出2条较粗的颏副神经。
6.外展神经起于延髓的前腹侧面、锥体前端两侧的斜方体后缘,分布于眼外侧直肌和眼球缩肌。
7.面神经起于延髓斜方体的前外侧部,在未伸出茎乳突孔之前发出岩大神经、镫骨肌神经、鼓索神经和一加入岩小神经的交通支,并接受一支来自前庭耳蜗神经的交通支和迷走神经的耳支。面神经穿出茎乳突孔后发出耳后神经、耳内支、腮腺支、二腹肌支、颈支、耳睑神经、颊背侧支和颊腹侧支。耳后神经和耳内支有侧支分布于腮腺和腮耳肌。颊背侧支在犬齿肌深面分为许多小支与眶下神经的分支形成一神经丛。颊腹侧支于下颌骨的外侧面与颏副神经有较粗的交通支相连。
8.前庭耳蜗神经连于延髓斜方体的背外侧缘,在内耳道门即分成上、下两支,即前庭部和耳蜗部。且在内耳道内,前庭耳蜗神经与面神经之间有2～3条细小分支相互吻合。
9.舌咽神经起于延髓外侧缘,其分支有鼓室神经、颈动脉窦支、茎突咽肌支、舌支、咽支、一与迷走神经耳支相连的交通支及一与面神经相连的交通支。
10.迷走神经起于延髓外侧缘,与舌咽神经和副神经相联合经颈静脉孔出颅腔。在颈前段分出耳支、喉前神经、咽喉食管气管总干和一些连于舌咽神经、舌下神经、第一颈神经和颈前神经节的交通支。
11.副神经起于中枢神经系统的脑神经根和脊神经根。脑神经根起于延髓侧缘;脊神经根起于脊髓前五颈节侧缘。延伸至鼓泡腹侧缘处在其内侧分出一较小的内侧支汇入迷走神经,其主干行经第一颈神经腹侧支时分为背、腹两支。背支沿途发出若干分支分布于臂头肌和肩胛横突肌,本干终止于斜方肌;腹支分支分布于枕—锁肌、胸下颌肌和胸乳突肌。
12.舌下神经起于延髓腹外侧面,穿过脑硬膜后汇集成一主干,经舌下神经管出颅腔。舌下神经穿过茎舌肌与舌骨舌肌间时,分出若干细小分支分布于颈舌肌、舌骨舌肌、颏舌骨肌与颏舌肌;其后,主干分成若干分支分布于舌骨舌肌、颏舌肌、颏舌骨肌和舌固有肌。舌下神经在延伸途中有交通支分别与迷走神经、副神经、颈前神经节和第一颈神经腹侧支相连。
13.头部交感神经
颈前神经节呈纺锤形,位于头长肌的前外侧面。其前部约25%被鼓泡覆盖,其余部分被茎突舌骨肌覆盖,且后缘与颈内动脉相平行。颈前神经节的前端发出5～6条颈内动脉神经,在头长肌的前外侧面伴随颈内动脉向前向背侧延伸约3～3.5cm后,围绕颈内动脉相互连接形成颈内动脉神经丛,该丛穿过鼓枕裂和岩枕裂到达颞骨磷部前缘,在此分出如下分支:岩深神经、颈动脉鼓室神经、海绵窦神经和颈静脉神经。其后端除发出较大的颈外动脉神经和交感干外,还发出数条通往舌咽神经、迷走神经及舌下神经。
14.头部副交感神经
14.1睫状神经节呈粟粒形,位于眼腹侧直肌后背侧和视神经腹外侧,与动眼神经腹侧支紧密相连(3/15标本的睫状神经节埋在动眼神经腹侧支内),接受来自鼻睫神经交通支,发出睫状短神经进入眼球。未见到副睫状神经节。
14.2耳神经节较大,呈倒三角形,位于腭帆张肌后上部外侧面。其后上端与岩小神经相连,前上端延续为一较粗的神经支加入颊神经。耳神经节内侧面光滑,外侧面大部分为上颌神经覆盖,且发出若干小支与之相连。耳神经节发出若干分支分布于鼓膜张肌、腭帆张肌和腭帆提肌。
14.3下颌神经节为一较大的呈梭形的神经节或为一组(5～6个)大小和形状不同的小神经节,位于舌下腺的外侧面、下颌舌骨肌和茎舌肌之间。接受来自舌神经的交通支,发出分支分布于下颌腺、舌下腺。
14.4翼腭神经节是大小和形状不同的一组神经节,一般相互融合或被神经纤维连为一体。在翼内侧肌背侧部的外侧面,位于翼腭神经丛。翼腭神经节发出若干分支分别汇入鼻后神经和腭大神经。
The origin, distribution, branch, shape and location of the cranial nerves in the head and neck of the yak were studied with gross anatomy method to explore the morphology mechanism of the cranial nerves of the yak to adapt to the ecological environment of the Qinghai-Tibetan plateau, which can provide the basic data for furthermore molecular and gene levels' research. The results are as followed:
1. The olfactory nerves arises from the mucous membrane of the caudal portions on the nasal septum and superior nasal meatus, and terminates in the rostroventral aspect of the olfactory bulb. The terminal nerve arises from the mucosa of the retro-superior part of the superior nasal meatus and blood vessel and glandular organ in the olfactory area epithelium of the retro-superior part of the nasal septum, and terminates in the medial surface of the olfactory trigon and subcallosal area. The vomeronasal nerve arises from the vomeronasal organ in the medial of the proximal wall of the anterior part of the nasal cavity and mucosa of the close together area and nasal septum, and ends in the olfactory gyrus of the anterio-medial margin of the olfactory triangle.
2. The optic nerve arises from the optic nerve disc of the retina of the eye bulb, enters the cranial cavity through the optic canal, runs retro-medially about 2～3 cm in the cranial cavity and terminates in the optic chiasm. It is circular cylinder shape and its diameter ranges from 4.60 mm to 5.50mm. The diameter of the optic nerve ranges from the optic chiasm to the eyeball about 0.50 mm. It runs like S-shape in the orbit, and receives one to two twigs from the ciliary ganglion during its course.
3. The oculomotor nerve origins from the intercrural fossa of the posterior part of the ventral aspect of the mesencephalic cerebral peduncle. Immediately, it divides into dorsal and ventral branches after leaving the cranial cavity through the foramen orbitorotundum. The two branches supply all bulbal muscles except the lateral rectus and dorsal oblique muscles. The ciliary nerv is from the ventral branch to associate with the ciliary ganglion.
4. The trochlear nerve comes from the canal between the colliculus caudalis of the quadrigeminal bodies (corpora quadrigemina) and rostral medullary velum, goes to the fossa orbitalis through the foramen orbitorotundum, and supplies the obliquus dorsalis muscle. It was connected the ophthalmic nerve with the communicating branches.
5. The trigeminal nerve arises from the dorsolateral side of the pons, and divides into the ophthalmic, maxillary and mandibular nerves.
The ophthalmic nerve gives off the zygomaticotemporal branch, lacrimal, frontal, frontal sinus nerves, muscular branches and nasociliary nerve. The nasociliary nerve divides into the infratrochlear and ethmoidal nerves. The ethmoidal nerve divides into the internal nasal and external nasal branches. There is a communicating branch in the nasociliary nerve and the ciliary ganglion.
The maxillary nerve gives off the zygomaticofacial, accessory zygomaticofacial branches, zygomatic bone, caudal nasal, greater palatine, lesser palatine and caudal superior alveolar nerves. The infraorbital nerve is the continuation of the maxillary nerve. The maxillary nerve is connected with the ophthalmic and buccal nerves by the major communicating branches. The caudal nasal nerve gives off some slender branches to the pterygopalatine plexus in the pterygopalatine fossa. The greater palatine nerve gives off accessory palatine nerve and collateral branches to enter the proximal wall of the nasal cavity.
The mandibular nerve gives off the deep temporal, masseteric, buccal, lingual, inferior alveolar, mylohyoid, medial peterygoid, later peterygoid and auriculotemporal nerves. The buccal nerve is connected with the maxillary, greater palatin nerves and dorsal buccal branch of the facial nerve by means of the communicating branches. The medial pterygoid muscle is also innervated by a branch of the buccal nerve. Two large accessory mental nerves arise from the inferior alveolar nerve.
6. The abducent nerve is originated from the posterior margin of the trapezoid body of the antero-ventral surface of the medulla oblongata and bilateral anterior extremity of the pyramid. The nerve branches supply to the rectus lateral and retractor bulbi muscles.
7. The facial nerve arises from the antero-lateral aspect of the trapezoid body of the medulla oblongata. Before its emergence through the stylomastiod foramen, the nerve gives off the greater petrosal, stapedial nerves, chorda tymapanic and communicating branches with the lesser petrosal nerve. It receives a communicating branch of the vestibulocochlear nerve and a auricular branch of the vagus nerve. After its emergence through the stylomastiod foramen, the nerve gives off the posterior auricular, auriculopalpebral nerves, internal auricular, parotid, digastric, cervical, dorsal buccal and ventral buccal branches. The caudal auricular nerve and the internal auricular branch give off wigs to the parotid gland and parotid auricularis muscle. The dorsal buccal branch divides into branches and forms a plexus with the branches of the infraorbital nerve under the canius muscle. The ventral buccal branch is connected with the accessory mental nerve on the lateral aspect of the mandible.
8. The vestibulocochlear nerve connects with the dorsolateral side of the trapezoid body of the medulla oblongata. In the internal acoustic meatus, it splites into the vestibular and cochlear parts, exchanging 2～3 twigs with the facial nerve.
9. The glossopharyngeal nerve arises from the lateral aspect of the medulla oblongata. It gives off the tympanic nerve, carotid sinus, stylopharyngeal, lingual, pharyngeal branch, a communiacating branch to join the auricular branch of the vagus and a communiacating branch to the facial nerve.
10. The vagus nerve arises from the lateral aspect of the medulla oblongata. It leaves the cranial cavity in company with the glossophoryngeal and accessory nerves through the jugular foramen. At the oranial end of the neck, the vagus nerve gives off the auricular branch, laryngis cranial nerve, the common trunk for the pharynx, larynx, oesophagus and trachea and some communicating branches which connect with the glossopharyngeal, hypoglossal nerves and cranial cervical ganglion.
11. The accessory nerve arises from the cranial spinal spinal roots of the central nervous system. The cranial root arises from the dorsolateral aspect of the medulla oblongata. The spinal root arises from the lateral aspect of fifth cervical segment of the spinal cord. When the nerve reaches to the ventral margin of the tympanic bulla, it gives off a thin internal branch, which joined to the vagus. And then, when the nerve crosses the ventral branch of the first cervical nerve, it divides into dorsal and ventral branches. The dorsal branch finally ramifies in the medial surface of the trapezius muscle, and it gives off some twigs to supply brachium cephalic and shoulder-blade processus transversus muscles. The vental branch gives off some branches to supply the cleido-occipitalis, sternomandibularis and sternomastoideus muscles.
12. The hypoglossal neve arises from the ventrolateral aspect of the medulla oblongata. The rootlets course laterally, perforate the dura mater and then unite in a single trunk. The trunk leaves the cranial cavity through the hypoglossal canal. When the hypoglossal nerve courses on the hypoglossal and digastricus muscles, it gives off several twigs to supply cervicum lingual, hyoglossus, geniohyoid and genioglossus muscles. And then, the trunk divides into several branches to supply hyoglossus, geniohyoid, genioglossus and lingua intrinsic muscles. At the coursing, some communicating branches connects with the vagus, accessory nerves, cranial cervical ganglion and the ventral branch of the first cervical nerve.
13. The sympathetic nerve in the head
The the cranial cervical ganglion is a greyish fusiform structure, which is located on the rostrolateral surface of the m. longus capitis. Roughly anterior 25% of the ganglion is covered by the tympanic bulla, the remainder by the m. stylohyoideus. The caudodorsal border of the ganglion is parallel to the internal carotid artery. Five or six internal carotid nerve branches arise from the rostrodorsal end of the cranial cervical ganglion and coursed rostrally parallel to each other about 3-3.5 cm along the rostrolateral surface of the m. longus capitis, following the internal carotid artery. Around the internal carotid artery, these branches forms the internal carotid plexus, which passed rostrally and dorsally through the tympano-occipital and petrooccipital fissures to gain the rostral border of the petrous temporal bone. There, the internal carotid plexus gives off four branches, deep petrosal, caroticotympanic, cavernous sinus and jugular nerves. The caudodorsal border of the cranial cervical ganglion gives off external carotid nerve and sympathetic trunk, together with some branches connecting the ganglion to the glossopharyngeal, vagus and hypoglossal nerves.
14. The parasympathetic nerve in the head
14.1 The ciliary ganglion is like a german millet, lies on the retrodorsal part of the ventral rectus of the eye and the ventrolateral part of the optic nerve and joins to the ventral branch of oculomotor nerve ( In the 3/15 specimens, the ciliary ganglion is hided in the ventral branch of oculomotor nerve) . It is connected with nasociliary nerves by communicative branches. The short ciliary nerve arises from the ciliary ganglion and enters the eyeball. But the yaks do have not the accessory ciliary ganglion.
14.2 The auricular ganglion is a triangle structure of the basilar part upward. It locates on the lateral aspect of the retro-superior part of the tensor veli palatini muscle. Its retro-superior extremity connects with the lesser petrosal nerve. The ventro-superior extremity give off a large branch to connect with the buccal nerve. The medialateral aspect is smooth and glossy. The large part of the lateral aspect is covered by the maxillary nerve and give off several twigs to join the maxillary nerve. The ganglioin give off several branches to supply tensor tympani, tensor veli palatini muscle and levator muscle of velum palatine.
14.3 The mandibular ganglion is a large fusiform ganglion, or is a lot of (five to six) small grayish structures. In the lateral aspect of the sublingual gland, it is located between the mylohyoid and cervicum lingual muscles. The ganglia are joined by a communicating branch from the lingual nerves and give off some branches to supply the mandibular and sublingual glands.
14.4 The pterygopalatine ganglia are a lot of different size and form ganglions, and frequently are fused or connected to one another by strands of the nerve fibers. In the lateral surface of the medial pterygoid muscle, they are located in the pterygopalatine nerve plexus. The ganglia send off several twigs to the caudal nasal and greater palatine nerves.
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