Auxin transport, metabolism, and signalling during nodule initiation: indeterminate and determinate nodules

Most legumes can form a unique type of lateral organ on their roots: root nodules. These structures host symbiotic nitrogen-fixing bacteria called rhizobia. Several different types of nodules can be found in nature, but the two best-studied types are called indeterminate and determinate nodules. These two types differ with respect to the presence or absence of a persistent nodule meristem, which consistently correlates with the cortical cell layers giving rise to the nodule primordia. Similar to other plant developmental processes, auxin signalling overlaps with the site of organ initiation and meristem activity. Here, we review how auxin contributes to early nodule development. We focus on changes in auxin transport, signalling, and metabolism during nodule initiation, describing both experimental evidence and computer modelling. We discuss how indeterminate and determinate nodules may differ in their mechanisms for generating localized auxin response maxima and highlight outstanding questions for future research

A Model for the Development of the Rhizobial and Arbuscular Mycorrhizal Symbioses in Legumes and Its Use to Understand the Roles of Ethylene in the Establishment of these two Symbioses

We propose a model depicting the development of nodulation and arbuscular mycorrhizae. Both processes are dissected into many steps, using Pisum sativum L. nodulation mutants as a guideline. For nodulation, we distinguish two main developmental programs, one epidermal and one cortical. Whereas Nod factors alone affect the cortical program, bacteria are required to trigger the epidermal events. We propose that the two programs of the rhizobial symbiosis evolved separately and that, over time, they came to function together. The distinction between these two programs does not exist for arbuscular mycorrhizae development despite events occurring in both root tissues. Mutations that affect both symbioses are restricted to the epidermal program. We propose here sites of action and potential roles for ethylene during the formation of the two symbioses with a specific hypothesis for nodule organogenesis. Assuming the epidermis does not make ethylene, the microsymbionts probably first encounter a regulatory level of ethylene at the epidermis–outermost cortical cell layer interface. Depending on the hormone concentrations there, infection will either progress or be blocked. In the former case, ethylene affects the cortex cytoskeleton, allowing reorganization that facilitates infection; in the latter case, ethylene acts on several enzymes that interfere with infection thread growth, causing it to abort. Throughout this review, the difficulty of generalizing the roles of ethylene is emphasized and numerous examples are given to demonstrate the diversity that exists in plants

23rd Century surprises: Long-term dynamics of the climate and carbon cycle under both high and net negative emissions scenarios

Future climate projections from Earth system models (ESMs) typically focus on the timescale of this century. We use a set of four ESMs and one Earth system model of intermediate complexity (EMIC) to explore the dynamics of the Earth’s climate and carbon cycles under contrasting emissions trajectories beyond this century, to the year 2300. The trajectories include a very high emissions, unmitigated fossil-fuel driven scenario, as well as a second mitigation scenario that diverges from the first scenario after 2040 and features an “overshoot”, followed by stabilization of atmospheric CO2 concentrations by means of large net-negative CO2 emissions. In both scenarios, and for all models considered here, the terrestrial system switches from being a net sink to either a neutral state or a net source of carbon, though for different reasons and centered in different geographic regions, depending on both the model and the scenario. The ocean carbon system remains a sink, albeit weakened by climate-carbon feedbacks, in all models under the high emissions scenario, and switches from sink to source in the overshoot scenario. The global mean temperature anomaly generally follows the trajectories of cumulative carbon emissions, except that 23rd-century warming continues after the cessation of carbon emissions in several models, both in the high emissions scenario and in one model in the overshoot scenario. While ocean carbon cycle responses qualitatively agree both in globally integrated and zonal-mean dynamics in both scenarios, the land models qualitatively disagree in zonal-mean dynamics, in the relative roles of vegetation and soil in driving C fluxes, in the response of the sink to CO2, and in the timing of the sink-source transition, particularly in the high emissions scenario. The lack of agreement among land models on the mechanisms and geographic patterns of carbon cycle feedbacks, alongside the potential for lagged physical climate dynamics to cause warming long after CO2 concentrations have stabilized, point to the possibility of surprises in the climate system beyond the 21st century time horizon, even under relatively mitigated global warming scenarios, which should be taken into consideration when setting global climate policy

In need of mediation: The relation between syntax and information structure

This paper defends the view that syntax does not directly interact with information structure. Rather, information structure affects prosody, and only the latter has an interface with syntax. We illustrate this point by discussing scrambling, focus preposing, and topicalization. The position entertained here implies that syntax is not very informative when one wants to narrow down the interpretation of terms such as “focus”, “topic”, etc

Role of the Arabidopsis PIN6 auxin transporter in auxin homeostasis and auxin-mediated development

Plant-specific PIN-formed (PIN) efflux transporters for the plant hormone auxin are required for tissue-specific directional auxin transport and cellular auxin homeostasis. The Arabidopsis PIN protein family has been shown to play important roles in developmental processes such as embryogenesis, organogenesis, vascular tissue differentiation, root meristem patterning and tropic growth. Here we analyzed roles of the less characterised Arabidopsis PIN6 auxin transporter. PIN6 is auxin-inducible and is expressed during multiple auxin–regulated developmental processes. Loss of pin6 function interfered with primary root growth and lateral root development. Misexpression of PIN6 affected auxin transport and interfered with auxin homeostasis in other growth processes such as shoot apical dominance, lateral root primordia development, adventitious root formation, root hair outgrowth and root waving. These changes in auxin-regulated growth correlated with a reduction in total auxin transport as well as with an altered activity of DR5-GUS auxin response reporter. Overall, the data indicate that PIN6 regulates auxin homeostasis during plant development.Christopher I. Cazzonelli, Marleen Vanstraelen, Sibu Simon, Kuide Yin, Ashley Carron-Arthur, Nazia Nisar, Gauri Tarle, Abby J. Cuttriss¤, Iain R. Searle, Eva Benkova, Ulrike Mathesius, Josette Masle, Jiří Friml, Barry J. Pogso