Stock D W
Department of Environmental, Population, and Organismic Biology, University of Colorado, Boulder, CO 80309-0334, USA.
Philos Trans R Soc Lond B Biol Sci. 2001 Oct 29;356(1414):1633-53. doi: 10.1098/rstb.2001.0917.
The construction of organisms from units that develop under semi-autonomous genetic control (modules) has been proposed to be an important component of their ability to undergo adaptive phenotypic evolution. The organization of the vertebrate dentition as a system of repeated parts provides an opportunity to study the extent to which phenotypic modules, identified by their evolutionary independence from other such units, are related to modularity in the genetic control of development. The evolutionary history of vertebrates provides numerous examples of both correlated and independent evolution of groups of teeth. The dentition itself appears to be a module of the dermal exoskeleton, from which it has long been under independent genetic control. Region-specific tooth loss has been a common trend in vertebrate evolution. Novel deployment of teeth and reacquisition of lost teeth have also occurred, although less frequently. Tooth shape differences within the dentition may be discontinuous (referred to as heterodonty) or graded. The occurrence of homeotic changes in tooth shape provides evidence for the decoupling of tooth shape and location in the course of evolution. Potential mechanisms for region-specific evolutionary tooth loss are suggested by a number of mouse gene knockouts and human genetic dental anomalies, as well as a comparison between fully-developed and rudimentary teeth in the dentition of rodents. These mechanisms include loss of a tooth-type-specific initiation signal, alterations of the relative strength of inductive and inhibitory signals acting at the time of tooth initiation and the overall reduction in levels of proteins required for the development of all teeth. Ectopic expression of tooth initiation signals provides a potential mechanism for the novel deployment or reacquisition of teeth; a single instance is known of a gene whose ectopic expression in transgenic mice can lead to ectopic teeth. Differences in shape between incisor and molar teeth in the mouse have been proposed to be controlled by the region-specific expression of signalling molecules in the oral epithelium. These molecules induce the expression of transcription factors in the underlying jaw mesenchyme that may act as selectors of tooth type. It is speculated that shifts in the expression domains of the epithelial signalling molecules might be responsible for homeotic changes in tooth shape. The observation that these molecules are regionally restricted in the chicken, whose ancestors were not heterodont, suggests that mammalian heterodonty may have evolved through the use of patterning mechanisms already acting on skeletal elements of the jaws. In general, genetic and morphological approaches identify similar types of modules in the dentition, but the data are not yet sufficient to identify exact correspondences. It is speculated that modularity may be achieved by gene expression differences between teeth or by differences in the time of their development, causing mutations to have cumulative effects on later-developing teeth. The mammalian dentition, for which virtually all of the available developmental genetic data have been collected, represents a small subset of the dental diversity present in vertebrates as a whole. In particular, teleost fishes may have a much more extensive dentition. Extension of research on the genetic control of tooth development to this and other vertebrate groups has great potential to further the understanding of modularity in the dentition.
从在半自主基因控制下发育的单元(模块)构建生物体,被认为是其进行适应性表型进化能力的一个重要组成部分。脊椎动物的牙列作为一个重复部分的系统,提供了一个机会来研究由其与其他此类单元的进化独立性所确定的表型模块,在多大程度上与发育的基因控制中的模块化相关。脊椎动物的进化史提供了许多牙齿群相关进化和独立进化的例子。牙列本身似乎是真皮外骨骼的一个模块,长期以来它一直处于独立的基因控制之下。特定区域的牙齿脱落是脊椎动物进化中的一个常见趋势。牙齿的新部署和失去牙齿后的重新获得也有发生,尽管频率较低。牙列内牙齿形状的差异可能是不连续的(称为异形齿)或渐变的。牙齿形状的同源异型变化的发生为牙齿形状和位置在进化过程中的解耦提供了证据。一些小鼠基因敲除、人类遗传性牙齿异常以及啮齿动物牙列中发育完全的牙齿与退化牙齿之间的比较,提示了特定区域进化性牙齿脱落的潜在机制。这些机制包括牙齿类型特异性起始信号的丧失、牙齿起始时作用的诱导和抑制信号相对强度的改变以及所有牙齿发育所需蛋白质水平的整体降低。牙齿起始信号的异位表达为牙齿的新部署或重新获得提供了一种潜在机制;已知有一个基因,其在转基因小鼠中的异位表达可导致异位牙齿。小鼠门齿和臼齿之间形状的差异被认为是由口腔上皮中信号分子的区域特异性表达所控制的。这些分子诱导下颌间充质中可能作为牙齿类型选择因子的转录因子的表达。据推测,上皮信号分子表达域的变化可能是牙齿形状同源异型变化的原因。观察到这些分子在鸡中是区域受限的,而鸡的祖先不是异形齿,这表明哺乳动物的异形齿可能是通过利用已经作用于颌骨骨骼元素的模式形成机制进化而来的。一般来说,遗传和形态学方法在牙列中识别出相似类型的模块,但数据还不足以确定确切的对应关系。据推测,模块化可能是通过牙齿之间的基因表达差异或其发育时间的差异来实现的,从而使突变对后期发育的牙齿产生累积影响。几乎所有可用的发育遗传学数据都是针对哺乳动物牙列收集的,而哺乳动物牙列只是整个脊椎动物中存在的牙齿多样性的一小部分。特别是,硬骨鱼类可能有更广泛的牙列。将牙齿发育的基因控制研究扩展到这个和其他脊椎动物群体,对于进一步理解牙列中的模块化具有巨大潜力。