本文選題：雄性生殖系統(tǒng) + 精子形成�。� 參考：《西北農(nóng)林科技大學(xué)》2016年博士論文

【摘要】：長翅目Mecoptera是全變態(tài)類昆蟲Holometabola中較古老的一個(gè)小類群,是唯一幼蟲期具有復(fù)眼的類群,在全變態(tài)類昆蟲系統(tǒng)發(fā)育研究中占據(jù)著獨(dú)特的地位。然而,長翅目的系統(tǒng)發(fā)育地位卻一直存在有爭議,亟需尋找更多的證據(jù)。精子是有性生殖動(dòng)物高度特化的雄配子。昆蟲精子具有快速、趨異進(jìn)化的特性,其形態(tài)、結(jié)構(gòu)(尤其是超微結(jié)構(gòu))復(fù)雜多樣,在不同類群間表現(xiàn)出顯著差異。精子形態(tài)的變化可為昆蟲分類、進(jìn)化及其系統(tǒng)發(fā)育分析和重建提供有價(jià)值的特征。但關(guān)于長翅目精子結(jié)構(gòu)的研究迄今仍然很少,僅涉及蝎蛉科Panorpidae、蚊蝎蛉科Bittacidae和雪蝎蛉科Boreidae等3科7種。而長翅目昆蟲的精子發(fā)生僅在研究蝎蛉科和蚊蝎蛉科精子超微結(jié)構(gòu)時(shí)對精子形成過程稍加描述,并未進(jìn)行詳細(xì)過程的研究。本文以蝎蛉科、蚊蝎蛉科、和擬蝎蛉科Panorpodidae昆蟲為研究對象,采用光學(xué)顯微鏡和電子顯微鏡技術(shù),研究了長翅目昆蟲的雄性生殖系統(tǒng)、精子發(fā)生以及精子超微結(jié)構(gòu),明確了精子發(fā)生的過程,比較了長翅目昆蟲不同類群間雄性生殖系統(tǒng)的差異以及精子超微結(jié)構(gòu),豐富了長翅目生殖生物學(xué)研究的內(nèi)容,并在精子結(jié)構(gòu)研究的基礎(chǔ)上,首次運(yùn)用精子超微結(jié)構(gòu)特征對長翅目進(jìn)行系統(tǒng)發(fā)育分析。主要研究結(jié)果如下:長翅目昆蟲雄性生殖系統(tǒng)主要包括1對對稱的精巢及兩個(gè)附睪,2條分區(qū)明顯的輸精管,1對特化的貯精囊,1對中胚層附腺,1個(gè)射精囊及其周圍的外胚層附腺(蝎蛉科新蝎蛉屬Neopanorpa及擬蝎蛉科除外)和陽莖。其中貯精囊特化為具有強(qiáng)烈分泌功能的器官,并不用于儲(chǔ)存精子。雄性生殖系統(tǒng)形態(tài)在長翅目科間變異較大,科內(nèi)變異較小,主要表現(xiàn)在精巢小管的數(shù)目,精巢和附睪的相對位置,囊后輸精管的長度,中胚層附腺的大小,以及外胚層附腺的有無等方面。首次詳細(xì)研究了長翅目昆蟲的精子發(fā)生。結(jié)果表明,長翅目昆蟲精子發(fā)生過程相似。精原細(xì)胞在精囊中以2n的方式,共經(jīng)歷7次連續(xù)分裂,產(chǎn)生128個(gè)精細(xì)胞。最后轉(zhuǎn)化形成的精子束中的精子通常低于128個(gè)。長翅目昆蟲精子各組成部分的形成大致如下:精子頂體由高爾基復(fù)合體產(chǎn)生的原頂體顆粒物質(zhì)形成;線形、致密的細(xì)胞核由圓形細(xì)胞核伸長及核內(nèi)染色質(zhì)凝集形成;軸絲由基體結(jié)構(gòu)產(chǎn)生;2條線粒體衍生物由大量分散的線粒體聚集、融合產(chǎn)生的副核轉(zhuǎn)化形成。精子形成過程中,細(xì)胞核核溝側(cè)旁出現(xiàn)2條側(cè)片結(jié)構(gòu),但該結(jié)構(gòu)在精子成熟后消失。此外,形成過程中在軸絲雙微管的外圍出現(xiàn)少量微管結(jié)構(gòu),但后者很快消失,使軸絲呈現(xiàn)9+2型微管結(jié)構(gòu)。研究發(fā)現(xiàn),長翅目昆蟲精子較長(1000μm以上),由較短的頭部、頸區(qū)和長鞭毛組成,且精子質(zhì)膜外具有一層縱向脊?fàn)畹亩嗵?蛋白質(zhì)鞘結(jié)構(gòu)。精子頭部包括兩部分,即由外層頂體囊和內(nèi)層頂體組成的雙層頂體,以及具有兩條側(cè)溝的細(xì)長細(xì)胞核。頸區(qū)由中心粒和中心粒側(cè)體組成;中心粒頂端通常插入細(xì)胞核底部,由9條雙微管組成;中心粒側(cè)體位于頸區(qū)最外圍,為致密的鞘狀結(jié)構(gòu)。鞭毛由1條軸絲、2條線粒體衍生物、1或2條副體和2條特殊的附結(jié)構(gòu)組成。其中2條線粒體衍生物大小不等,較小的1條隨鞭毛的延伸而消失,較大的1條占據(jù)后半部鞭毛的大部分體積。副體較短,來源于中心粒側(cè)體物質(zhì)。2條附結(jié)構(gòu)細(xì)小,橫切多呈三角形,延伸至鞭毛末端。鞭毛末端線粒體衍生物逐漸減小,軸絲結(jié)構(gòu)也因缺失雙微管的動(dòng)力蛋白臂和輻射鏈結(jié)構(gòu)而變得混亂無序,直至消失。研究發(fā)現(xiàn),長翅目精子超微結(jié)構(gòu)科間變異較大,科內(nèi)變異較小�？崎g變異主要表現(xiàn)在細(xì)胞核形狀、核溝尺寸、軸絲結(jié)構(gòu)、副體數(shù)目、以及較大線粒體衍生物等。本研究首次發(fā)現(xiàn)擬蝎蛉科精子軸絲為9+9+2型結(jié)構(gòu)。我們推測,蝎蛉科和蚊蝎蛉科中出現(xiàn)的9+2型軸絲,很可能是由9+9+2型進(jìn)化逆轉(zhuǎn)而來的。本文基于精子超微結(jié)構(gòu),利用最大簡約法構(gòu)建了系統(tǒng)發(fā)育樹,進(jìn)行了具泵總目Antliophora的系統(tǒng)發(fā)育分析,這是首次利用精子超微結(jié)構(gòu)對長翅目進(jìn)行系統(tǒng)發(fā)育分析。結(jié)果表明,長翅目為單系群,與蚤目Siphonaptera互為姊妹群關(guān)系;長翅目內(nèi)擬蝎蛉科與蝎蛉科互為姊妹群關(guān)系,而非蚊蝎蛉科。
[Abstract]:The long winged Mecoptera is a relatively old group of all metamorphosis insect Holometabola, and it is the only group with compound eyes in the only larval stage. It occupies a unique position in the phylogenetic study of all metamorphosis insects. However, the phylogenetic status of long wings has always been controversial, and it is urgent to find more evidence. Sperm is sexual. The highly specialized male gamete of colonies. Insect sperm has the characteristics of rapid and heterogeneous evolution, its morphology, structure (especially ultramicrostructure) is complex and diverse, showing significant differences among different groups. The changes in sperm morphology can provide valuable characteristics for the classification of insects, evolution, phylogenetic analysis and reconstruction. The study of substructure is still rare, involving only 3 families, 7 species, such as Panorpidae, mosquitididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididididido, and the spermatogenesis of the long winged insects are only slightly described in the study of spermatogenesis in the spermatozoa of the family scorpiididididaand mosquitoes. The male reproductive system, spermatogenesis and sperm ultrastructure of the long winged insects were studied by optical microscopy and electron microscopy. The differences in male reproductive system between different groups of long winged insects and spermatogenesis were compared and the spermatogenesis of Panorpodidae were compared. The ultrastructure of the substructure enriches the research of the reproductive biology of the long dteroptera. On the basis of the study of the sperm structure, the phylogenetic analysis of the long wterptera is used for the first time. The main results are as follows: the male reproductive system of the long winged insects mainly consists of 1 pairs of symmetrical sperms and two epididymis, 2 distinct divisions. The vas deferens, 1 pairs of seminal vesicles, 1 pairs of mesoderm glands, 1 eminal vesicles and the surrounding ectodermal attachment (except for the genus Neopanorpa and pheididido) and Yang stems. The seminal vesicles are specially secreted as organs that have strong secretory functions and are not used for storage of sperm. The variation of Kone is small, mainly in the number of the seminiferous tubules, the relative position of the spermary and epididymis, the length of the spermatogonial duct, the size of the mesoderm glands, and the presence of the ectoderm gland. The spermatogenesis of the long winged insects is studied in detail for the first time. The results show that the spermatogenesis of the long winged insects is similar. In the seminal vesicle, a total of 7 continuous divisions and 128 spermatozoa are produced in a total of 7 successive divisions. The sperm in the spermatozoa is usually lower than 128. The formation of the sperm components of the long winged insects is as follows: the sperm acrosome formed by the PRODIC particles produced by the Golgi complex; the linear, compact nuclei are round and thin. The nucleus is elongated and the chromatin agglutination is formed in the nucleus; the shaft filament is produced by the matrix structure, and the 2 mitochondrial derivatives are formed by a large number of dispersed mitochondria and the paramuclear fusion produced by the fusion. In the process of sperm formation, 2 side segments appear on the side of the nucleus furrow, but the structure disappears after the spermatozoon is mature. In addition, in the formation process, the axis wire is double. A small number of microtubules appear on the periphery of the microtubule, but the latter quickly disappears, making the shaft filament present 9+2 microtubule structure. It is found that the sperm of the long winged insect is longer (more than 1000 mu m), consisting of a shorter head, the neck region and the long flagellum, and the sperm plasma membrane has a longitudinal ridge like polysaccharide protein sheath structure. The sperm head consists of two parts, that is, the sperm head consists of the sperm head. The bilayer acrosome consisting of the outer apical sac and the inner apical body and the elongated nuclei with two side trenches. The neck region is composed of central particles and central granular bodies; the tip of the central kernel usually inserts the bottom of the nucleus and consists of 9 double microtubules; the central granulocyte is located at the periphery of the neck region and is a dense sheath. The flagellum consists of 1 axes and 2 lines. Body derivatives, 1 or 2 pairs of accessory bodies and 2 special attached structures, of which 2 mitochondrial derivatives vary in size, the smaller 1 strips disappear with the extension of the flagellum, and the larger 1 strips occupy most of the volume of the posterior half of the flagellum. The side body is short, and the.2 strips from the central granular substance are small in structure, and in the transverse section, extending to the end of flagellum. The mitochondrial derivative at the end of the flagellum gradually decreased, and the structure of the axis filament became chaotic and disordered by the deletion of the two microtubule's brachial arm and the structure of the radiation chain. It was found that the ultrastructural variations of the spermatozoa were larger and the Kone variation was smaller. The interfamily variation mainly manifested in the shape of the nucleus, the size of the nuclear furrow, the structure of the axis silk. The number of side bodies, as well as the larger mitochondrial derivatives. For the first time, the sperm axis of the paramedicinic family was found to be a 9+9+2 type structure. We speculated that the 9+2 type axis in the scorpion and mosquitoes may be reversed by the evolution of the 9+9+2 type. The phylogenetic analysis of the Antliophora was performed. This is the first time to analyze the phylogenetic analysis of the long wterptera using sperm ultrastructure. The results show that the long winged group is a monophyletic group and a sister group with the flea Siphonaptera, and the pteroptera is a sister group relationship with the family scorpiidido, not the mosquito.
【學(xué)位授予單位】：西北農(nóng)林科技大學(xué)
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2016
【分類號(hào)】：Q964
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本文編號(hào)：1949773