植物叶缘形态的发育调控机理

doi:10.17520/biods.2018127

摘要/Abstract

摘要：

生物多样性研究的关键问题之一是表型多样性的形成和演化机制, 因为表型多样性与物种多样性密切相关, 同时又承载着遗传和环境的变异信息。植物的叶具有丰富的形态多样性, 而叶形多样性很大程度上体现在叶边缘形态的变异。叶边缘的形态可从全缘、锯齿状到具有不同程度(深浅)和不同式样(羽状或掌状、回数等)的裂片(在发育研究中复叶的小叶也描述为裂片)。关于叶缘齿/裂的发育调控机制, 在拟南芥(Arabidopsis thaliana)、碎米荠(Cardamine hirsuta)、番茄(Solanum lycopersicum)等模式植物中已有较深入的探讨。研究发现, 很多转录因子、小分子RNA及植物激素对叶齿/裂或小叶的形成具有调控作用, 其中生长素输出途径中的转录因子NAM/CUC、miR164以及高浓度生长素的反馈调控可能起到核心作用, 而且该调控模块在真双子叶植物中较为保守; TCP类、SPL类转录因子和其他一些miRNA也在生长素输出途径中发挥作用; 关于KNOX家族转录因子的作用, 虽然多数研究是围绕复叶的形态建成, 但也有数据显示其在叶裂发育中发挥作用。此外, 对拟南芥和碎米荠等十字花科植物的研究还发现, 调控基因RCO通过抑制小叶/裂片之间的细胞增殖而对小叶/叶裂的发育发挥作用。本文综述上述多角度的研究进展, 并尝试概括叶边缘形态的发育调控网络, 为关于叶缘形态多样性形成机制的研究提供可参考的切入点。

关键词: 叶缘齿/裂, 形态发生, 演化发育, 调控网络

Abstract

The mechanism of formation and evolution of phenotypic diversity is one of the key problems in biodiversity science because phenotype diversity is not only a marker of species diversity, but carries the designs adjusted to environments. Plant leaves exhibit a great deal of morphological variation. Such variation is attributed largely to changes of leaf marginal architecture. Leaf marginal shapes can be described as entire, serrate, lobed (varying in depth and patterns) and dissected (also referred to as compound leaf). The molecular mechanism controlling the development of leaf marginal shape has been intensively studied in Arabidopsis thaliana, Cardamine hirsuta, Solanum lycopersicum, and some other plants. Many important regulatory factors such as transcription factors, small RNAs and plant hormones have been found involved in the development of leaf serration or dissection. Among those factors, the transcription factor NAM/CUC, miR164 and auxin in the auxin efflux module play a central role through a feedback loop, and this regulatory module appears to be conserved across the eudicots; the transcription factors TCPs, SPLs and some other miRNAs also take part in the auxin efflux pathway. Transcription factors of the KNOX family play roles in the development of leaf lobes as well although most of the researches about KNOX genes have focused on their regulation of the morphogenesis of compound leaves. In addition, studies in Arabidopsis, Cardamine and other taxa of the Brassicaceae have shown that the gene RCO promotes the development of leaf dissection by repressing growth at the flanks of protrusions generated by CUC-auxin patterning. The present paper reviews the recent progress and integrate the major results of researches about the molecular mechanisms that underlie leaf serration or dissection. We hope this may provide reference for unraveling the morphogenetic origin of the spectacular diversity of leaf marginal shapes.

Key words: leaf serration/dissection, morphogenesis, evolutionary development, regulatory network

柯锦秀, 陈多, 郭延平 (2018) 植物叶缘形态的发育调控机理. 生物多样性, 26, 988-997. DOI: 10.17520/biods.2018127.

Jinxiu Ke, Duo Chen, Yanping Guo (2018) Designing leaf marginal shapes: Regulatory mechanisms of leaf serration or dissection. Biodiversity Science, 26, 988-997. DOI: 10.17520/biods.2018127.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2018127

https://www.biodiversity-science.net/CN/Y2018/V26/I9/988

图/表 2

图1 叶缘锯齿发育的调控模型(综合拟南芥和碎米荠中的研究, 基于Barkoulas等(2007)和Runions和Tsiantis (2017)两文中的图修订而成)。该调控模型的核心是转录因子CUC2 (红色)与生长素(黄色)之间的反馈作用环: CUC2指导PIN1蛋白在细胞中重新定位, 将生长素从两个方向逆浓度梯度运到一个汇聚点; 生长素的积累又反过来通过激活miR164抑制CUC2的表达。CUC2和miR164在共同区域活动, 相互平衡, 调控着生长素的极性运输, 生长素促进叶齿的生长。另外, 其他生长调节因子也参与修饰叶齿/裂的形态发育, 比如: 碎米荠等十字花科植物的RCO类基因抑制叶齿两侧细胞的生长, 调控更多的齿/裂形态式样。

Fig. 1 A model for the molecular regulation of the development of leaf marginal serration according to studies on Arabidopsis thaliana and Cardamine hirsute (Barkoulas et al, 2007; Runions & Tsiantis, 2017). At the heart of the model is a feedback loop between CUC2 and auxin activities. CUC2 is required for PIN1-mediated auxin polar transport (horrow arrows); in turn, auxin activity maxima at the tip of the developing serration activates miR164 which represses CUC2 posttranscriptionally and generates an interspersed pattern of auxin maxima and CUC2 expression at the leaf margin. MIR164 and CUC2 are expressed in partially overlapping regions at the sinus of the serrations. Auxin enhances outgrowth of the serrations. There are additional growth regulators modulating leaf growth to shape the form of protrusions, for instance, RCO inhibits growth in indentations, producing more dissected forms.

图2 叶边缘形态发育调控网络图, 示各因子间的相互作用。实线表示已有大量研究清晰阐明的相互作用; 虚线则表示相关研究报道较少, 尚不太明确的相互作用; 橙色箭头表示促进叶齿/裂表型的形成。

Fig. 2 The molecular regulating network underlining the development leaf serration/dissection. The solid lines indicate the relatively clear interactions between molecules, while the dotted lines suggest unsure interactions reported by a few studies. The orange arrows suggest promotion of leaf serration/dissection.

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