蝎蛉科昆虫胚胎消化道形成与幼虫腹足同源性研究
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摘要
利用扫描电子显微镜和光学显微镜,对长翅目Mecoptera蝎蛉科Panorpidae昆虫的胚胎发育过程进行组织学和外部形态研究。以太白蝎蛉Panorpa obtusa为材料,研究了蝎蛉胚胎期的消化道形成,证实口道和肛道均由外胚层发育而来,而中肠由内胚层发育而来;马氏管由肛道前端形成,因此起源于外胚层。并且以大蝎蛉Panorpa magna为材料,研究了蝎蛉胚胎期腹足的形成及其超微结构,观察了蝎蛉腹足的发育过程,讨论了腹足与胸足间的非同源性关系。本研究丰富了长翅目胚胎学知识,为进一步深入探讨昆虫胚胎发育机理奠定了基础。
结果表明,太白蝎蛉的口道起源于头叶中部的浅凹陷,肛道起源于胚胎腹部末端,几乎与口道同时出现。与口道相比,肛道发育速度很快,肠道伸长并最终形成一个环状。证明了口道和肛道都是由外胚层发育而来。六瓣状的贲门瓣出现在口道的末端,并逐渐浸入发育中的中肠前端。在马氏管开口处紧紧相连着具有瓣状褶皱结构的幽门瓣。
六条马氏管由肛道前端两侧的突起发育而来,每侧的突起各发育出三个马氏管,分别向各个方向朝胚胎前端发散延长。伸达胚胎第二腹节时,马氏管折回并继续发育延伸至第八腹节。因此证实了马氏管也起源于外胚层。
研究发现,中肠由处于胚胎两极的前中肠韧和后中肠韧共同发育而来,前中肠韧和后中肠韧细胞分别在胚胎腹面发育形成两条细胞带,相向生长至胚胎中部第二腹节与第三腹节间结合形成中肠原基。观察了中肠原基各部分细胞的不同形态以及在胚胎背合后中肠上皮细胞的主要特点,同时比较了口道、中肠和后肠的形态差异。作为前肠和中肠的分界,贲门瓣与中肠之间有一层单细胞组成的封闭膜结构。类似的膜结构也出现于中肠与幽门瓣之间。这种膜结构在孵化前几小时内消失。
通过对大蝎蛉胸足和腹足组织切片和外部形态对比,发现蝎蛉的腹足并非真正的附肢,而是在真附肢的内侧新产生形成的构造。腹部的附肢在胚胎发育后期停止发育,并在幼虫期留下比较明显的痕迹,因此证实长翅目幼虫的腹足与胸足之间并非同源构造。
The morphogenesis during embryogenesis of the family Panorpidae in Mecoptera was studied by using scanning electron microscopy and light microscopy. Based on the embryo of Panorpa obtusa, we investigated the formation of the alimentary canal. The results show that the stomodaeum and proctodaeum are developed from ectoderm and midgut is derived from endoderm. The Malpighian tubules are formed by anterior part of proctodaeum, therefore they are also developed from the ectoderm. In addition, we studied the formation and ultrastructure of the abdominal proleg in Panorpa magna during embryogenesis. The process of prolegs development was observed and they are confirmed to not to be homologous with the thoracic legs. The present study gathers new data for the mecopteran embryology and lays a sound foundation for the further research of the developmental mechanism in insects.
From our study, we found that the furrow of the stomodaeum is derived from the middle part of the protocephalic lobe. The proctodaeum originates from the caudal end of the embryonic abdomen, which almost occurs simultaneously with the stomodaeum. It grows faster than the stomodaeum and eventually elongates to form a loop. This shows that both the stomodaeum and proctodaeum arise from the ectoderm. Six ruga-shaped cardiac valves appear at the posterior end of stomodaeum, gradually embedding to the front of the developing midgut. The pylorus valves were observed to have layers of valve-like ruga just behind the openings of the Malpighian tubules.
Six Malpighian tubules are derived from the tubers which has three Malpighian tubules respectively on the anterior proctodaeal wall, extending forwards to various orientations. When the Malpighian tubules reach the second abdominal segment, they turn back until reaching the eighth abdominal segment. It is evident that they are also ectodermal in origin.
The midgut arises from the anterior and posterior midgut rudiments. The two cell-bands are formed at the posterior end of the stomodaeum and the anterior end of the proctodaeum respectively. These two bands grow and elongate towards each other along the venter of the embryo between the yolk and mesodermal cells, eventually meet and connect each other at between the second and third abdominal segments. The cell morphology varies among different regions of the midgut rudiment and the main characteristics of the midgut epithelial cells after the dorsal closure of the embryo development are described. Morphological differentiation among the stomodaeum, midgut and proctodaeum were observed. Between the cardiac valves and the rudimental midgut there are some monolayered cells forming a membrane as the border between stomodaeum and midgut rudiment. The same membrane structure is formed between the rudimental midgut and the pylorus valves. Both the anterior and the posterior cellular membranes begin to degenerate just a few hours before hatching.
Compared with the thoracic legs in histology and external morphology, we confirm that the abdominal prologs are not true appendages, but newly formed structure on the mesial side of the true appendages which are homologous with thoracic legs. During the later embryogenesis, the true abdominal appendages stop developing and become reduced gradually, their remnants could still be observed in larval stage. Therefore our conclusion is that the prolegs of Panorpa could not be serially homologous with the thoracic legs in Mecoptera.
结果表明,太白蝎蛉的口道起源于头叶中部的浅凹陷,肛道起源于胚胎腹部末端,几乎与口道同时出现。与口道相比,肛道发育速度很快,肠道伸长并最终形成一个环状。证明了口道和肛道都是由外胚层发育而来。六瓣状的贲门瓣出现在口道的末端,并逐渐浸入发育中的中肠前端。在马氏管开口处紧紧相连着具有瓣状褶皱结构的幽门瓣。
六条马氏管由肛道前端两侧的突起发育而来,每侧的突起各发育出三个马氏管,分别向各个方向朝胚胎前端发散延长。伸达胚胎第二腹节时,马氏管折回并继续发育延伸至第八腹节。因此证实了马氏管也起源于外胚层。
研究发现,中肠由处于胚胎两极的前中肠韧和后中肠韧共同发育而来,前中肠韧和后中肠韧细胞分别在胚胎腹面发育形成两条细胞带,相向生长至胚胎中部第二腹节与第三腹节间结合形成中肠原基。观察了中肠原基各部分细胞的不同形态以及在胚胎背合后中肠上皮细胞的主要特点,同时比较了口道、中肠和后肠的形态差异。作为前肠和中肠的分界,贲门瓣与中肠之间有一层单细胞组成的封闭膜结构。类似的膜结构也出现于中肠与幽门瓣之间。这种膜结构在孵化前几小时内消失。
通过对大蝎蛉胸足和腹足组织切片和外部形态对比,发现蝎蛉的腹足并非真正的附肢,而是在真附肢的内侧新产生形成的构造。腹部的附肢在胚胎发育后期停止发育,并在幼虫期留下比较明显的痕迹,因此证实长翅目幼虫的腹足与胸足之间并非同源构造。
The morphogenesis during embryogenesis of the family Panorpidae in Mecoptera was studied by using scanning electron microscopy and light microscopy. Based on the embryo of Panorpa obtusa, we investigated the formation of the alimentary canal. The results show that the stomodaeum and proctodaeum are developed from ectoderm and midgut is derived from endoderm. The Malpighian tubules are formed by anterior part of proctodaeum, therefore they are also developed from the ectoderm. In addition, we studied the formation and ultrastructure of the abdominal proleg in Panorpa magna during embryogenesis. The process of prolegs development was observed and they are confirmed to not to be homologous with the thoracic legs. The present study gathers new data for the mecopteran embryology and lays a sound foundation for the further research of the developmental mechanism in insects.
From our study, we found that the furrow of the stomodaeum is derived from the middle part of the protocephalic lobe. The proctodaeum originates from the caudal end of the embryonic abdomen, which almost occurs simultaneously with the stomodaeum. It grows faster than the stomodaeum and eventually elongates to form a loop. This shows that both the stomodaeum and proctodaeum arise from the ectoderm. Six ruga-shaped cardiac valves appear at the posterior end of stomodaeum, gradually embedding to the front of the developing midgut. The pylorus valves were observed to have layers of valve-like ruga just behind the openings of the Malpighian tubules.
Six Malpighian tubules are derived from the tubers which has three Malpighian tubules respectively on the anterior proctodaeal wall, extending forwards to various orientations. When the Malpighian tubules reach the second abdominal segment, they turn back until reaching the eighth abdominal segment. It is evident that they are also ectodermal in origin.
The midgut arises from the anterior and posterior midgut rudiments. The two cell-bands are formed at the posterior end of the stomodaeum and the anterior end of the proctodaeum respectively. These two bands grow and elongate towards each other along the venter of the embryo between the yolk and mesodermal cells, eventually meet and connect each other at between the second and third abdominal segments. The cell morphology varies among different regions of the midgut rudiment and the main characteristics of the midgut epithelial cells after the dorsal closure of the embryo development are described. Morphological differentiation among the stomodaeum, midgut and proctodaeum were observed. Between the cardiac valves and the rudimental midgut there are some monolayered cells forming a membrane as the border between stomodaeum and midgut rudiment. The same membrane structure is formed between the rudimental midgut and the pylorus valves. Both the anterior and the posterior cellular membranes begin to degenerate just a few hours before hatching.
Compared with the thoracic legs in histology and external morphology, we confirm that the abdominal prologs are not true appendages, but newly formed structure on the mesial side of the true appendages which are homologous with thoracic legs. During the later embryogenesis, the true abdominal appendages stop developing and become reduced gradually, their remnants could still be observed in larval stage. Therefore our conclusion is that the prolegs of Panorpa could not be serially homologous with the thoracic legs in Mecoptera.
引文
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