羊膜卵的系统发育和进化史,Journal of Morphology

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羊膜卵的系统发育和进化史
Journal of Morphology ( IF 1.5 ) Pub Date : 2021-05-15 , DOI: 10.1002/jmor.21380
J Matthias Starck ₁ , James R Stewart _{1,

2} , Daniel G Blackburn ₃

Affiliation

我们回顾了比较系统发育框架中羊膜卵和胚胎的形态特征，包括现存脊椎动物的所有主要进化枝。我们讨论了与脊椎动物卵进化历史分析相关的 40 个特征。特别注意细胞卵黄囊、蛋壳和胚外膜的形态。许多通常被赋予羊膜动物的特征，例如大卵黄囊、延迟的卵沉积和陆生繁殖，已经在众多脊椎动物进化枝中独立和集中进化。我们使用系统发育特征图谱和祖先特征状态重建作为工具来识别形态进化的序列、顺序和模式，并推断出羊膜卵进化史的假设。除了羊膜和尿囊绒毛膜外，羊膜动物祖先还拥有交配器官（在大多数鸟类中次要减少）、内部受精和包含原始条纹或早期体节阶段胚胎的卵的延迟沉积。除了羊膜、尿囊绒毛膜和羊膜类型的蛋壳外，这些特征在几乎所有主要的水生脊椎动物进化枝中都集中进化，可能是为了响应选择性因素，例如卵捕食、卵发育的恶劣环境条件，或调整幼鸟的孵化到有利的季节。羊膜的一个重要功能特征是其生肌收缩力，可移动（早期）胚胎并防止生长中的胚胎粘附在胚胎外材料上。羊膜和充满液体的羊膜腔的这种功能可能是在含有发育中胚胎的卵子延迟沉积的要求下进化而来的。尿囊绒毛膜是支持胚胎发育的临时胚胎交换器官。一种可能的进化场景是，羊膜卵呈现出一种进化，为陆地繁殖铺平了进化途径。正如许多来自羊膜动物的例子所表明的那样，脊椎动物在陆地上进行了多次繁殖——羊膜卵提供了一种“解决方案”，使征服土地进行繁殖成为可能。一种可能的进化场景是，羊膜卵呈现出一种进化，为陆地繁殖铺平了进化途径。正如许多来自羊膜动物的例子所表明的那样，脊椎动物在陆地上进行了多次繁殖——羊膜卵提供了一种“解决方案”，使征服土地进行繁殖成为可能。一种可能的进化场景是，羊膜卵呈现出一种进化，为陆地繁殖铺平了进化途径。正如许多来自羊膜动物的例子所表明的那样，脊椎动物在陆地上进行了多次繁殖——羊膜卵提供了一种“解决方案”，使征服土地进行繁殖成为可能。

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Phylogeny and evolutionary history of the amniote egg

We review morphological features of the amniote egg and embryos in a comparative phylogenetic framework, including all major clades of extant vertebrates. We discuss 40 characters that are relevant for an analysis of the evolutionary history of the vertebrate egg. Special attention is given to the morphology of the cellular yolk sac, the eggshell, and extraembryonic membranes. Many features that are typically assigned to amniotes, such as a large yolk sac, delayed egg deposition, and terrestrial reproduction have evolved independently and convergently in numerous clades of vertebrates. We use phylogenetic character mapping and ancestral character state reconstruction as tools to recognize sequence, order, and patterns of morphological evolution and deduce a hypothesis of the evolutionary history of the amniote egg. Besides amnion and chorioallantois, amniotes ancestrally possess copulatory organs (secondarily reduced in most birds), internal fertilization, and delayed deposition of eggs that contain an embryo in the primitive streak or early somite stage. Except for the amnion, chorioallantois, and amniote type of eggshell, these features evolved convergently in almost all major clades of aquatic vertebrates possibly in response to selective factors such as egg predation, hostile environmental conditions for egg development, or to adjust hatching of young to favorable season. A functionally important feature of the amnion membrane is its myogenic contractility that moves the (early) embryo and prevents adhering of the growing embryo to extraembryonic materials. This function of the amnion membrane and the liquid-filled amnion cavity may have evolved under the requirements of delayed deposition of eggs that contain developing embryos. The chorioallantois is a temporary embryonic exchange organ that supports embryonic development. A possible evolutionary scenario is that the amniote egg presents an exaptation that paved the evolutionary pathway for reproduction on land. As shown by numerous examples from anamniotes, reproduction on land has occurred multiple times among vertebrates—the amniote egg presenting one “solution” that enabled the conquest of land for reproduction.

更新日期：2021-06-13

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