Too many faces for TOO MANY MOUTHS?
Too many faces for TOO MANY MOUTHS?
复制标题
DOI：
10.1111/nph.13827
复制
发表时间：
2016-05-01
期刊：
影响因子：
9.4
通讯作者：
Mena, Montana
中科院分区：
文献类型：
--
作者：
de Marcos, Alberto;Trivino, Magdalena;Mena, Montana
几十年来，气孔生理学一直吸引着植物科学家。然而，直到最近，他们的注意力才转向研究这些结构如何从原皮层细胞分化，以及成熟叶片和其他气生器官的功能空间模式如何出现。我们的理解气孔发育有很大的进步，在过去的20年，由于采用拟南芥作为模式物种。第一个参与气孔发育的基因是在A. Yang & Sack（1995）在通过失去正确气孔间距的突变体的叶表皮表型的正向遗传筛选中，其中一个突变体产生了大量的气孔聚集在引人注目的集群。这个基因座的功能丧失导致了这种表型，因此它配得上一个醒目的名字：太多的嘴（TMM）。Yang & Sack（1995）还鉴定了产生成对气孔的第二个突变体，并将该位点命名为FOUR LIPS。由Fred Sack的实验室进行的这项开创性工作所建立的命名法在气孔发育领域已经普及，并且从那时起鉴定的大多数基因都获得了响亮的名字，强调气孔作为植物的“嘴”的观点。thaliana，基于先前在其他物种中的工作（Paliwal，1967; Sachs & Novoplansky，1993），从而建立了该过程的当前观点（参见支持信息图S1）。原胚层细胞获得分生组织样母细胞（MMC）的身份，并不对称分裂，产生一个小细胞，分生组织样细胞，其表现为干细胞，并成为气孔细胞谱系的创始人。多达三个不对称的分裂，在一个向内的螺旋方向，将分生组织放置在由较大的姐妹细胞形成的结构（气孔复合体）的中心，这些姐妹细胞最终将分化为铺路细胞或成为MMC，启动卫星气孔谱系。最终，分生组织停止分裂并转变为保卫母细胞，进行最终的对称分裂，其结果是保卫细胞对分化形成气孔。在该谱系中的各种细胞中，分生组织和保卫母细胞统称为气孔前体细胞。基于tmm子叶和叶表型，Yang & Sack（1995）提出TMM限制分生组织的产生，阻止其产生。
Stomatal physiology has attracted plant scientists for many decades. However, it is only recently that their attention has turned to the study of how these structures differentiate from the protodermal cells, and how functional spatial patterns in mature leaves and other aerial organs emerge. Our understanding of stomatal development has advanced enormously in the last 20 yr, thanks to the adoption of Arabidopsis thaliana as a model species. The first genes involved in stomatal development were identified in A. thaliana by Yang & Sack (1995), in a forward genetic screen through the leaf epidermal phenotype of mutants that had lost the correct stomatal spacing. One of these mutants produced overabundant stomata grouped in striking clusters. The locus whose loss of function produced this phenotype thus deserved the striking name given to it: TOO MANY MOUTHS (TMM). Yang & Sack (1995) also identified a second mutant that produced paired stomata and named the locus FOUR LIPS. The nomenclature established by this seminal work, undertaken by Fred Sack’s laboratory, has been pervasive in the field of stomatal development, and most of the genes identified since then have received sonorous names stressing the view of the stomata as ‘mouths’ of the plant.In their pioneering work, Sack and his colleague described the course of stomatal development in A. thaliana, based on prior work in other species (Paliwal, 1967; Sachs & Novoplansky, 1993), and thus established the current view of the process (see Supporting Information Fig. S1). A protodermal cell acquires the identity of a meristemoid mother cell (MMC) and divides asymmetrically, producing a small cell, the meristemoid, which behaves as a stem cell and becomes the founder of a stomatal cell lineage. Up to three asymmetric divisions, oriented in an inward spiral, place the meristemoid at the centre of a structure (the stomatal complex) formed by the larger sister cells, which will eventually differentiate as pavement cells or become MMCs, initiating a satellite stomatal lineage. Eventually, the meristemoid stops dividing and transits to a guard mother cell, committed to a final symmetric division whose result is the guard cell pair that differentiate to form the stoma. Among the various cells in the lineage, meristemoids and guard mother cells are collectively referred to as stomatal precursor cells. On the basis of the tmm cotyledon and leaf phenotype, Yang & Sack (1995) proposed that TMM restricts meristemoid production, prevents its