Stewart James R, Heulin Benoit, Surget-Groba Yann
Department of Biological Sciences, East Tennessee State University, Johnson City, TN 37614-1710, USA.
Zoology (Jena). 2004;107(4):289-314. doi: 10.1016/j.zool.2004.07.004. Epub 2004 Nov 2.
Reproductive mode has been remarkably labile among squamate reptiles and the evolutionary transition from oviparity to viviparity commonly has been accompanied by a shift in the pattern of embryonic nutrition. Structural specializations for placental transfer of nutrients during intrauterine gestation are highly diverse and many features of the extraembryonic membranes of viviparous species differ markedly from those of oviparous species. However, because of a high degree of evolutionary divergence between the species used for comparisons it is likely that the observed differences arose secondarily to the evolution of viviparity. We studied development of the extraembryonic membranes and placentation in the reproductively bimodal lizard Lacerta vivipara because the influence of reproductive mode on the structural/functional relationship between mothers and embryos can best be understood by studying the most recent evolutionary events. Lecithotrophic viviparity has evolved recently within this species and, although populations with different reproductive modes are allopatric, oviparous and viviparous forms interbreed in the laboratory and share many life history characteristics. In contrast to prior comparisons between oviparous and viviparous species, we found no differences in ontogeny or structure of the extraembryonic membranes between populations with different reproductive modes within L. vivipara. However, we did confirm conclusions from previous studies that the tertiary envelope of the egg, the eggshell, is much reduced in the viviparous population. These conclusions support a widely accepted model for the evolution of squamate placentation. We also found support for work published nearly 80 years ago that the pattern of development of the yolk sac of L. vivipara is unusual and that a function of a unique structure of squamate development, the yolk cleft, is hematopoiesis. The structure of the yolk sac splanchnopleure of L. vivipara is inconsistent with a commonly accepted model for amniote yolk sac function and we suggest that a long standing hypothesis that cells from the yolk cleft participate in yolk digestion requires further study.
在有鳞类爬行动物中,生殖模式一直非常不稳定,从卵生到胎生的进化转变通常伴随着胚胎营养模式的转变。子宫内妊娠期间胎盘营养物质转运的结构特化高度多样,胎生物种的胚外膜的许多特征与卵生物种明显不同。然而,由于用于比较的物种之间存在高度的进化差异,观察到的差异很可能是在胎生进化之后次生出现的。我们研究了生殖方式双峰型蜥蜴胎生蜥蜴(Lacerta vivipara)胚外膜的发育和胎盘形成,因为通过研究最近的进化事件,能最好地理解生殖模式对母体与胚胎之间结构/功能关系的影响。卵黄营养型胎生最近在该物种中进化而来,尽管具有不同生殖模式的种群是异域分布的,但卵生和胎生形式在实验室中可以杂交,并且具有许多共同的生活史特征。与之前对卵生和胎生物种的比较不同,我们发现在胎生蜥蜴不同生殖模式的种群之间,胚外膜的个体发育或结构没有差异。然而,我们确实证实了先前研究的结论,即卵的第三层包膜——蛋壳,在胎生种群中大大减少。这些结论支持了一个被广泛接受的有鳞类胎盘形成进化模型。我们还支持了近80年前发表的研究成果,即胎生蜥蜴的卵黄囊发育模式不寻常,并且有鳞类发育的独特结构——卵黄裂的一个功能是造血。胎生蜥蜴卵黄囊脏壁层的结构与一个被广泛接受的羊膜动物卵黄囊功能模型不一致,我们认为一个长期存在的假设,即来自卵黄裂的细胞参与卵黄消化,需要进一步研究。