Auxin and embryo axis formation: the ends in sight?

The major axis of polarity of the plant embryo serves as a reference for the formation of meristems and, thus, for all subsequent development. Mechanisms underlying the establishment of the embryo axis itself have remained elusive. This is now changing with recent reports documenting a role for auxin in embryo axis formation. Auxin accumulates dynamically at specific positions that correlate with developmental decisions in early embryogenesis, and this ties developmental decisions to both transport regulators and components of the response machinery. A major challenge for the future is to determine how auxin-dependent processes interact with other as yet unknown factors to mediate differential gene expression patterns in early embryogenesis.     

Is there a role for auxin in early embryogenesis?

The very young embryo of a flowering plant is not an idealsystem in which to study the effects of auxin. Conversely, auxin isusually not considered as a major component of developmental processesin early embryogenesis. However, recent findings from both experimentalstudies in Brassica and analyses of developmental mutants inArabidopsis make it worthwhile to examine critically thepossibility that auxin may have a role in early embryogenesis. In thisreview, we will focus on specific processes, such as formation of anapical-basal axis of polarity and the initiation of the primary rootmeristem. To provide a conceptual framework in which to discuss possibleeffects of auxin, we will first briefly summarise essential features ofearly embryogenesis in Arabidopsis. This will be followed by anevaluation of relevant data suggesting a role for auxin in axisformation and root meristem initiation. Finally, we will discuss a fewexperimental approaches that we believe are necessary to examine whetheror not auxin plays a role in fundamental processes of earlyembryogenesis.     

生长素在植物胚胎早期发育中的作用

有性生殖是有花植物的一个重要特征, 胚胎则是实现有性生殖和世代交替的重要载体。植物胚胎从双受精开始, 经历了合子极性建立、顶基轴形成、细胞层分化和器官形成等过程, 这些过程都受到生长素的调控。近年来的研究表明, 生长素在生物合成、极性运输和信号转导3个层面上调控胚胎的发育过程。该文以双子叶植物拟南芥(Arabidopsis thaliana)为例, 综述了生长素对胚胎早期发育过程, 包括合子极性和顶基轴建立、表皮原特化和对称模式转变、胚根原特化和根尖分生组织形成及茎尖分生组织形成等发育的调控机制。     

The influence of heat stress on auxin distribution in transgenic B. napus microspores and microspore-derived embryos

Plant embryogenesis is regulated by differential distribution of the plant hormone auxin. However, the cells establishing these gradients during microspore embryogenesis remain to be identified. For the first time, we describe, using the DR5 or DR5rev reporter gene systems, the GFP- and GUS-based auxin biosensors to monitor auxin during Brassica napus androgenesis at cellular resolution in the initial stages. Our study provides evidence that the distribution of auxin changes during embryo development and depends on the temperature-inducible in vitro culture conditions. For this, microspores (mcs) were induced to embryogenesis by heat treatment and then subjected to genetic modification via Agrobacterium tumefaciens. The duration of high temperature treatment had a significant influence on auxin distribution in isolated and in vitro-cultured microspores and on microspore-derived embryo development. In the “mild” heat-treated (1 day at 32 °C) mcs, auxin localized in a polar way already at the uni-nucleate microspore, which was critical for the initiation of embryos with suspensor-like structure. Assuming a mean mcs radius of 20 μm, endogenous auxin content in a single cell corresponded to concentration of 1.01 μM. In mcs subjected to a prolonged heat (5 days at 32 °C), although auxin concentration increased dozen times, auxin polarization was set up at a few-celled pro-embryos without suspensor. Those embryos were enclosed in the outer wall called the exine. The exine rupture was accompanied by the auxin gradient polarization. Relative quantitative estimation of auxin, using time-lapse imaging, revealed that primordia possess up to 1.3-fold higher amounts than those found in the root apices of transgenic MDEs in the presence of exogenous auxin. Our results show, for the first time, which concentration of endogenous auxin coincides with the first cell division and how the high temperature interplays with auxin, by what affects delay early establishing microspore polarity. Moreover, we present how the local auxin accumulation demonstrates the apical–basal axis formation of the androgenic embryo and directs the axiality of the adult haploid plant.     

Taking the very first steps: from polarity to axial domains in the early Arabidopsis embryo

Arabidopsis embryos follow a predictable sequence of cell divisions, facilitating a genetic analysis of their early development. Both asymmetric divisions and cell-to-cell communication are probably involved in generating specific gene expression domains along the main axis within the first few division cycles. The function of these domains is not always understood, but recent work suggests that they may serve as a basis for organizing polar auxin flux. Auxin acts as systemic signal throughout the life cycle and, in the embryo, has been demonstrated to direct formation of the main axis and root initiation at the globular stage. At about the same time, root versus shoot fates are imposed on the incipient meristems by the expression of antagonistic regulators at opposite poles of the embryo. Some of the key features of the embryonic patterning process have emerged over the past few years and may provide the elements of a coherent conceptual framework.     

Auxin Control of Embryo Patterning

Plants start their life as a single cell, which, during the process of embryogenesis, is transformed into a mature embryo with all organs necessary to support further growth and development. Therefore, each basic cell type is first specified in the early embryo, making this stage of development excellently suited to study mechanisms of coordinated cell specification—pattern formation. In recent years, it has emerged that the plant hormone auxin plays a prominent role in embryo development. Most pattern formation steps in the early Arabidopsis embryo depend on auxin biosynthesis, transport, and response. In this article, we describe those embryo patterning steps that involve auxin activity, and we review recent data that shed light on the molecular mechanisms of auxin action during this phase of plant development.     

Abscisic Acid Represses Growth of the Arabidopsis Embryonic Axis after Germination by Enhancing Auxin Signaling

Under unfavorable environmental conditions, the stress phytohormone ABA inhibits the developmental transition from an embryo in a dry seed into a young seedling. We developed a genetic screen to isolate Arabidopsis thaliana mutants whose early seedling development is resistant to ABA. Here, we report the identification of a recessive mutation in AUXIN RESISTANT1 (AUX1), encoding a cellular auxin influx carrier. Although auxin is a major morphogenesis hormone in plants, little is known about ABA–auxin interactions during early seedling growth. We show that aux1 and pin2 mutants are insensitive to ABA-dependent repression of embryonic axis (hypocotyl and radicle) elongation. Genetic and physiological experiments show that this involves auxin transport to the embryonic axis elongation zone, where ABA enhances the activity of an auxin-responsive promoter. We propose that ABA represses embryonic axis elongation by potentiating auxin signaling in its elongation zone. This involves repression of the AUXIN INDUCIBLE (Aux/IAA) gene AXR2/IAA7, encoding a key component of ABA- and auxin-dependent responses during postgerminative growth.     

Polarity,Continuity, and Alignment in Plant Vascular Strands

Plant vascular cells are joined end to end along uninterrupted lines to connect shoot organs with roots; vascular strands are thus polar, continuous, and internally aligned. What controls the formation of vascular strands with these properties? The “auxin canalization hypothesis”—based on positive feedback between auxin flow through a cell and the cell's capacity for auxin transport—predicts the selection of continuous files of cells that transport auxin polarly, thus accounting for the polarity and continuity of vascular strands. By contrast, polar, continuous auxin transport—though required—is insufficient to promote internal alignment of vascular strands, implicating additional factors. The auxin canalization hypothesis was derived from the response of mature tissue to auxin application but is consistent with molecular and cellular events in embryo axis formation and shoot organ development. Objections to the hypothesis have been raised based on vascular organizations in callus tissue and shoot organs but seem unsupported by available evidence. Other objections call instead for further research; yet the inductive and orienting influence of auxin on continuous vascular differentiation remains unique.     

High auxin stimulates callus through SDG8-mediated histone H3K36 methylation in Arabidopsis

Callus induction,which results in fate transition in plant cells,is considered as the first and key step for plant regeneration.This process can be stimulated in different tissues by a callus-inducing medium(CIM),which contains a high concentration of phytohormone auxin.Although a few key regulators for callus induction have been identified,the multiple aspects of the regulatory mechanism driven by high levels of auxin still need further investigation.Here,we find that high auxin induces callus ...