TOR acts as a metabolic gatekeeper for auxin-dependent lateral root initiation in Arabidopsis thaliana.

Plant organogenesis requires matching the available metabolic resources to developmental programs. In Arabidopsis, the root system is determined by primary root-derived lateral roots (LRs), and adventitious roots (ARs) formed from non-root organs. Lateral root formation entails the auxin-dependent activation of transcription factors ARF7, ARF19, and LBD16. Adventitious root formation relies on LBD16 activation by auxin and WOX11. The allocation of shoot-derived sugar to the roots influences branching, but how its availability is sensed for LRs formation remains unknown. We combine metabolic profiling with cell-specific interference to show that LRs switch to glycolysis and consume carbohydrates. The target-of-rapamycin (TOR) kinase is activated in the lateral root domain. Interfering with TOR kinase blocks LR initiation while promoting AR formation. The target-of-rapamycin inhibition marginally affects the auxin-induced transcriptional response of the pericycle but attenuates the translation of ARF19, ARF7, and LBD16. TOR inhibition induces WOX11 transcription in these cells, yet no root branching occurs as TOR controls LBD16 translation. TOR is a central gatekeeper for root branching that integrates local auxin-dependent pathways with systemic metabolic signals, modulating the translation of auxin-induced genes.

SNF1-RELATED KINASE 1 and TARGET OF RAPAMYCIN control light-responsive splicing events and developmental characteristics in etiolated Arabidopsis seedlings.

The kinases SNF1-RELATED KINASE 1 (SnRK1) and TARGET OF RAPAMYCIN (TOR) are central sensors of the energy status, linking this information via diverse regulatory mechanisms to plant development and stress responses. Despite the well-studied functions of SnRK1 and TOR under conditions of limited or ample energy availability, respectively, little is known about the extent to which the 2 sensor systems function and how they are integrated in the same molecular process or physiological context. Here, we demonstrate that both SnRK1 and TOR are required for proper skotomorphogenesis in etiolated Arabidopsis (Arabidopsis thaliana) seedlings, light-induced cotyledon opening, and regular development in light. Furthermore, we identify SnRK1 and TOR as signaling components acting upstream of light- and sugar-regulated alternative splicing events, expanding the known action spectra for these 2 key players in energy signaling. Our findings imply that concurring SnRK1 and TOR activities are required throughout various phases of plant development. Based on the current knowledge and our findings, we hypothesize that turning points in the activities of these sensor kinases, as expected to occur upon illumination of etiolated seedlings, instead of signaling thresholds reflecting the nutritional status may modulate developmental programs in response to altered energy availability.

Plant cell surface immune receptors-Novel insights into function and evolution.

Plants use surface resident and intracellular immune receptors to provide robust immunity against microbial infections. The contribution of the two receptor types to plant immunity differs spatially and temporally. The ongoing identification of new plant cell surface immune receptors and their microbial-derived immunogenic ligands reveal a previously unexpected complexity of plant surface sensors involved in the detection of specific microbial species. Comparative analyses of the plant species distribution of cell surface immune receptors indicate that plants harbor larger sets of genus- or species-specific surface receptors in addition to very few widespread pattern sensors. Leucine-rich repeat surface and intracellular immune sensors emerge as two polymorphic receptor classes whose evolutionary trajectories appear to be linked. This is consistent with their functional cooperativity in providing full plant immunity.

From signals to stem cells and back again.

During plant development, organ morphology and body architecture are dynamically adjusted in response to a changing environment. This developmental plasticity is based on precisely controlled maintenance of primary, as well as programmed initiation of pluripotent stem cell populations during secondary- and de novo meristem formation (reviewed in [1-3]). Plant stem cells are found exclusively in specific locations that are defined by relative position within the growing tissue. It follows that stem cell fate is primarily instructed by endogenous signals that dynamically define the stem cell niche in response to tissue topography [4]. Furthermore, plant stem cell activity is strongly dependent on developmental stage, suggesting that they are sensitive to long range signaling from distant organs, including the root [5,6••]. And finally, environmental signals exert a major influence allowing plants to cope with the plethora of highly variable environmental parameters during their life-cycle [7]. Integrating tissue level positional information with long range developmental cues, as well as environmental signals requires intricate molecular mechanisms that allow to filter, classify, and balance diverse inputs and translate them into appropriate local cell behavior. In this short review, we aim to highlight advances in identifying the relevant signals, their mode of action, as well as the mechanisms of information processing in stem cells of the shoot apical meristem (SAM).

Integration of light and metabolic signals for stem cell activation at the shoot apical meristem.

A major feature of embryogenesis is the specification of stem cell systems, but in contrast to the situation in most animals, plant stem cells remain quiescent until the postembryonic phase of development. Here, we dissect how light and metabolic signals are integrated to overcome stem cell dormancy at the shoot apical meristem. We show on the one hand that light is able to activate expression of the stem cell inducer WUSCHEL independently of photosynthesis and that this likely involves inter-regional cytokinin signaling. Metabolic signals, on the other hand, are transduced to the meristem through activation of the TARGET OF RAPAMYCIN (TOR) kinase. Surprisingly, TOR is also required for light signal dependent stem cell activation. Thus, the TOR kinase acts as a central integrator of light and metabolic signals and a key regulator of stem cell activation at the shoot apex.