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整合发育进化模式与机制:以腹足纲齿舌为例的案例研究

INTEGRATING DEVELOPMENTAL EVOLUTIONARY PATTERNS AND MECHANISMS: A CASE STUDY USING THE GASTROPOD RADULA.

作者信息

Guralnick R P, Lindberg D R

机构信息

Department of Integrative Biology and Museum of Paleontology, University of California, Berkeley, California, 94720-4780.

出版信息

Evolution. 1999 Apr;53(2):447-459. doi: 10.1111/j.1558-5646.1999.tb03780.x.

Abstract

Determining the connection between ontogeny and phylogeny continues to be a major theme in biology. However, few studies have combined dissection of pattern and process that lead to transformation of complex morphological structures. Here we examine the patterns and processes of shape change in a model system-the gastropod radula. This system is a simple one having only two processes: initial secretion and postsecretional movement of teeth. However, it produces a tremendous amount of shape variability and fusion patterns. To determine both pattern and mechanism of shape change in an evolutionary context, we use three complementary approaches and datasets. First, we use a phylogenetic hypothesis to determine the polarity of developmental events. Second, we perform a morphometric analysis of shape change using relative warp analysis that allows us to locate and compare the direction and magnitude of ontogenetic and phylogenetic shape divergence. These comparisons are the basis for testing hyptheses of heterochrony and heterotopy, and we show how our results do not conform to expectations of pure heterochrony. The rejection of heterochrony as a hypothesis is based on empirically demonstrating (1) initial shape differs in each taxon; (2) a single dimension of shape variability does not simultaneously describe ontogenetic and evolutionary shape changes; and (3) a significantly different shape and size covariance between taxa. This rejection is probably based on spatial changes in initial conditions and not spatial changes caused by the process itself. Finally, we construct a mechanistic model that explains how shape change happens based on the sequence of events during ontogeny. By using the parameters in the model as characters in the phylogenetic dataset, we show that different parts of the system have arisen at different times and become co-opted into the process. By integrating our analyses together we show that spatial process parameters can be responsible for our nonspatial patterns and that different ontogenetic processes can create similar end morphologies.

摘要

确定个体发育与系统发育之间的联系一直是生物学的一个主要主题。然而,很少有研究将导致复杂形态结构转变的模式和过程剖析结合起来。在这里,我们研究了一个模型系统——腹足纲齿舌——形状变化的模式和过程。这个系统很简单,只有两个过程:牙齿的初始分泌和分泌后的移动。然而,它产生了大量的形状变异性和融合模式。为了在进化背景下确定形状变化的模式和机制,我们使用了三种互补的方法和数据集。首先,我们使用系统发育假设来确定发育事件的极性。其次,我们使用相对扭曲分析对形状变化进行形态测量分析,这使我们能够定位和比较个体发育和系统发育形状差异的方向和大小。这些比较是检验异时性和异位性假设的基础,并且我们展示了我们的结果如何不符合纯异时性的预期。作为一种假设,异时性被否定是基于经验证明:(1)每个分类单元的初始形状不同;(2)形状变异性的单一维度不能同时描述个体发育和进化形状变化;(3)分类单元之间形状和大小协方差存在显著差异。这种否定可能基于初始条件的空间变化,而不是过程本身引起的空间变化。最后,我们构建了一个机制模型,该模型解释了个体发育过程中基于事件序列的形状变化是如何发生的。通过将模型中的参数用作系统发育数据集中的特征,我们表明系统的不同部分在不同时间出现,并被纳入这个过程。通过将我们的分析整合在一起,我们表明空间过程参数可以解释我们的非空间模式,并且不同的个体发育过程可以产生相似的最终形态。

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