Plant postembryonic development takes place in the meristems, where stem cells self-renew and produce daughter cells that differentiate and give rise to different organ structures. For the maintenance of meristems, the rate of differentiation of daughter cells must equal the generation of new cells: How this is achieved is a central question in plant development. In the Arabidopsis root meristem, stem cells surround a small group of organizing cells, the quiescent center. Together they form a stem cell niche [1, 2], whose position and activity depends on the combinatorial action of two sets of genes - PLETHORA1 (PLT1) and PLETHORA2 (PLT2)[3, 4] and SCARECROW (SCR) and SHORTROOT (SHR)[2] - as well as on polar auxin transport. In contrast, the mechanisms controlling meristematic cell differentiation remain unclear. Here, we report that cytokinins control the rate of meristematic cell differentiation and thus determine root-meristem size via a two-component receptor histidine kinase-transcription factor signaling pathway. Analysis of the root meristems of cytokinin mutants, spatial cytokinin depletion, and exogenous cytokinin application indicates that cytokinins act in a restricted region of the root meristem, where they antagonize a non-cell-autonomous cell-division signal, and we provide evidence that this signal is auxin.

Plant growth and development are sustained by meristems. Meristem activity is controlled by auxin and cytokinin, two hormones whose interactions in determining a specific developmental output are still poorly understood. By means of a comprehensive genetic and molecular analysis in Arabidopsis, we show that a primary cytokinin-response transcription factor, ARR1, activates the gene SHY2/IAA3 (SHY2), a repressor of auxin signaling that negatively regulates the PIN auxin transport facilitator genes: thereby, cytokinin causes auxin redistribution, prompting cell differentiation. Conversely, auxin mediates degradation of the SHY2 protein, sustaining PIN activities and cell division. Thus, the cell differentiation and division balance necessary for controlling root meristem size and root growth is the result of the interaction between cytokinin and auxin through a simple regulatory circuit converging on the SHY2 gene.

Biotic and abiotic stresses limit agricultural yields, and plants are often simultaneously exposed to multiple stresses. Combinations of stresses such as heat and drought or cold and high light intensity have profound effects on crop performance and yields. Thus, delineation of the regulatory networks and metabolic pathways responding to single and multiple concurrent stresses is required for breeding and engineering crop stress tolerance. Many studies have described transcriptome changes in response to single stresses. However, exposure of plants to a combination of stress factors may require agonistic or antagonistic responses or responses potentially unrelated to responses to the corresponding single stresses. To analyze such responses, we initially compared transcriptome changes in 10 Arabidopsis (Arabidopsis thaliana) ecotypes using cold, heat, high-light, salt, and flagellin treatments as single stress factors as well as their double combinations. This revealed that some 61% of the transcriptome changes in response to double stresses were not predictable from the responses to single stress treatments. It also showed that plants prioritized between potentially antagonistic responses for only 5% to 10% of the responding transcripts. This indicates that plants have evolved to cope with combinations of stresses and, therefore, may be bred to endure them. In addition, using a subset of this data from the Columbia and Landsberg erecta ecotypes, we have delineated coexpression network modules responding to single and combined stresses.