Plants interact with diverse microbes including those that result in nutrient-acquiring symbioses. In order to balance the energy cost with the benefit gained, plants employ a systemic negative feedback loop to control the formation of these symbioses. This is particularly well-understood in nodulation, the symbiosis between legumes and nitrogen-fixing rhizobia, and is known as autoregulation of nodulation (AON). However, much less is understood about the autoregulation of the ancient arbuscular mycorrhizal symbioses that form between Glomeromycota fungi and the majority of land plants. Elegant physiological studies in legumes have indicated there is at least some overlap in the genes and signals that regulate these two symbioses but there are major gaps in our understanding. In this paper we examine the hypothesis that the autoregulation of mycorrhizae (AOM) pathway shares some elements with AON but that there are also some important differences. By reviewing the current knowledge of the AON pathway, we have identified important directions for future AOM studies. We also provide the first genetic evidence that CLV2 (an important element of the AON pathway) influences mycorrhizal development in a non-legume, tomato and review the interaction of the autoregulation pathway with plant hormones and nutrient status. Finally, we discuss whether autoregulation may play a role in the relationships plants form with other microbes.
Plants form mutualistic nutrient acquiring symbioses with microbes, including arbuscular mycorrhizal fungi. The formation of these symbioses is costly and plants employ a negative feedback loop termed autoregulation of mycorrhizae (AOM) to limit arbuscular mycorrhizae (AM) formation. We provide evidence for the role of one leucine-rich-repeat receptor like kinase (FAB), a hydroxyproline O-arabinosyltransferase enzyme (FIN) and additional evidence for one receptor like protein (SlCLV2) in the negative regulation of AM formation in tomato. Reciprocal grafting experiments suggest that the FAB gene acts locally in the root, while the SlCLV2 gene may act in both the root and the shoot. External nutrients including phosphate and nitrate can also strongly suppress AM formation. We found that FAB and FIN are required for nitrate suppression of AM but are not required for the powerful suppression of AM colonisation by phosphate. This parallels some of the roles of legume homologs in the autoregulation of the more recently evolved symbioses with nitrogen-fixing bacteria leading to nodulation. This deep homology in the symbiotic role of these genes suggests that in addition to the early signalling events that lead to the establishment of AM and nodulation, the autoregulation pathway might also be considered part of the common symbiotic toolkit that enabled plants to form beneficial symbioses.
This paper furthers our understanding of lock-on that is induced by periodic external forcing. The effect of forcing phase difference is investigated. An extended linear theory is proposed to predict the centers of various lock-on regimes, including harmonic, subharmonic, and superharmonic lock-on, in a parametric map spanned by the forcing frequency and phase difference. It reveals that when the forcing frequency is equal to the natural vortex shedding frequency or its integer multiple, harmonic or subharmonic lock-on occurs at particular forcing phase differences, whereas when the forcing frequency is a submultiple of the natural shedding frequency, superharmonic lock-on occurs. To confirm this theory and also further determine the shape and size of each lock-on regime, a series of numerical simulations is conducted on a circular-cylinder flow system with periodic external forcing being realized by a pair of synthetic jets (SJs). At a Reynolds number 100 and under moderate SJ forcing, five lock-on regimes are captured, including the primary, secondary, tertiary, and first-and second-superharmonic lock-on. It is found that these lock-on regimes are generally in a rhomboidal shape, and their size gradually reduces when the SJ frequency is away from the natural vortex shedding frequency. With these simulations, the aerodynamic forces and wake formation in each lock-on regime are analyzed and compared, with the discussion being focused on the effects of SJ frequency and phase difference. Furthermore, stability analysis is conducted to reveal more flow physics related to lock-on.
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