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SUMMARY Strigolactones (SLs) are key regulators of shoot growth and responses to environmental stimuli. Numerous studies have indicated that nitrogen (N) limitation induces SL biosynthesis, suggesting that SLs may play a pivotal role in coordinating systemic responses to N availability, but this idea has not been clearly demonstrated. Here, we generated triple knockout mutants in the SL synthesis gene TaDWARF17 (TaD17) in bread wheat and investigated their phenotypic and transcriptional responses under N limitation, aiming to elucidate the role of SLs in the adaptation to N limitation. Tad17 mutants display typical SL mutant phenotypes, and fail to adapt their shoot growth appropriately to N. Despite exhibiting an increased tillering phenotype, Tad17 mutants continued to respond to N limitation by reducing tiller number, suggesting that SLs are not the sole regulators of tillering in response to N availability. RNA‐seq analysis of basal nodes revealed that the loss of D17 significantly altered the transcriptional response of N‐responsive genes, including changes in the expression profiles of key N response master regulators. Crucially, our findings suggest that SLs are required for the transcriptional downregulation of cytokinin (CK) synthesis and signalling in response to N limitation. Collectively, our results suggest that SLs are essential for the appropriate morphological and transcriptional adaptation to N limitation in wheat, and that the repressive effect of SLs on shoot growth is partly mediated by their repression of CK synthesis. [ABSTRACT FROM AUTHOR]

The rhizosphere is a complex physical and chemical interface between plants and their underground environment, both biotic and abiotic. Plants exude a large number of chemicals into the rhizosphere in order to manipulate these biotic and abiotic components. Among such chemicals are strigolactones, ancient signalling molecules that in flowering plants act as both internal hormones and external rhizosphere signals. Plants exude strigolactones to communicate with their preferred symbiotic partners and neighbouring plants, but at least some classes of parasitic organisms are able to 'crack' these private messages and eavesdrop on the signals. In this review, we examine the intentional consequences of strigolactone exudation, and also the unintentional consequences caused by eavesdroppers. We examine the molecular mechanisms by which strigolactones act within the rhizosphere, and attempt to understand the enigma of the strigolactone molecular diversity synthesized and exuded into the rhizosphere by plants. We conclude by looking at the prospects of using improved understanding of strigolactones in agricultural contexts. [ABSTRACT FROM AUTHOR]

Vascular plants, unlike bryophytes, have a strong root-shoot dichotomy in which the tissue systems are mutually interdependent; roots are completely dependent on shoots for photosynthetic sugars, and shoots are completely dependent on roots for water and mineral nutrients. Long-distance communication between shoot and root is therefore critical for the growth, development and survival of vascular plants, especially with regard to variable environmental conditions. However, this long-distance signalling does not appear an ancestral feature of land plants, and has likely arisen in vascular plants to service the radical alterations in body-plan seen in this taxon. In this review, we examine the defined hormonal root-to-shoot and shoot-to-root signalling pathways that coordinate the growth of vascular plants, with a particular view to understanding how these pathways may have evolved. We highlight the completely divergent roles of isopentenyl-adenine and trans -zeatin cytokinin species in long-distance signalling, and ask whether cytokinin can really be considered as a single class of hormones in the light of recent research. We also discuss the puzzlingly sparse evidence for auxin as a shoot-to-root signal, the evolutionary re-purposing of strigolactones and gibberellins as hormonal signals, and speculate on the possible role of sugars as long-distance signals. We conclude by discussing the 'design principles' of long-distance signalling in vascular plants. [ABSTRACT FROM AUTHOR]

Summary: Strigolactones are an important class of plant signalling molecule with both external rhizospheric and internal hormonal functions in flowering plants. The past decade has seen staggering progress in strigolactone biology, permitting highly detailed understanding of their signalling, synthesis and biological roles – or so it seems. However, phylogenetic analyses show that strigolactone signalling mediated by the D14‐SCFMAX2‐SMXL7 complex is only one of a number of closely related signalling pathways, and is much less ubiquitous in land plants than might be expected. The existence of closely related pathways, such as the KAI2‐SMAX1 module, challenges many of our assumptions about strigolactones, and in particular emphasises how little we understand about the specificity of strigolactone signalling with respect to related signalling pathways. In this review, we examine recent advances in strigolactone signalling, taking a holistic evolutionary view to identify the ambiguities and uncertainties in our understanding. We highlight that while we now have highly detailed molecular models for the core mechanism of D14‐SMXL7 signalling, we still do not understand the ligand specificity of D14, the specificity of its interaction with SMXL7, nor the specificity of SMXL7 function. Our analysis therefore identifies key areas requiring further study. [ABSTRACT FROM AUTHOR]

The shoot systems of plants are built by the action of the primary shoot apical meristem, established during embryogenesis. In the axil of each leaf produced by the primary meristem, secondary axillary shoot apical meristems are established. The dynamic regulation of the activity of these axillary meristems gives shoot systems their extraordinary plasticity of form. The ability of plants to activate or repress these axillary meristems appropriately requires communication between meristems that is environmentally sensitive. The transport network of the plant hormone auxin has long been implicated as a central player in this tuneable communication system, with other systemically mobile hormones, such as strigolactone and cytokinin, acting in part by modulating auxin transport. Until recently, the polar auxin transport stream, which provides a high conductance auxin transport route down stems dominated by the auxin export protein PIN-FORMED1 (PIN1), has been the focus for understanding long range auxin transport in the shoot. However, recently additional auxin exporters with important roles in the shoot have been identified, including PIN3, PIN4 and PIN7. These proteins contribute to a wider less polar stem auxin transport regime, which we have termed connective auxin transport (CAT), because of its role in communication across the shoot system. Here we present a genetic analysis of the role of CAT in shoot branching. We demonstrate that in Arabidopsis, CAT plays an important role in strigolactone-mediated shoot branching control, with the triple pin3pin4pin7 mutant able to suppress partially the highly branched phenotype of strigolactone deficient mutants. In contrast, the branchy phenotype of mutants lacking the axillary meristem-expressed transcription factor, BRANCHED1 (BRC1) is unaffected by pin3pin4pin7. We further demonstrate that mutation in the ABCB19 auxin export protein, which like PIN3 PIN4 and PIN7 is widely expressed in stems, has very different effects, implicating ABCB19 in auxin loading at axillary bud apices. [ABSTRACT FROM AUTHOR]

Background: Strigolactones (SLs) are a class of plant hormones that control many aspects of plant growth. The SL signalling mechanism is homologous to that of karrikins (KARs), smoke-derived compounds that stimulate seed germination. In angiosperms, the SL receptor is an α/β-hydrolase known as DWARF14 (D14); its close homologue, KARRIKIN INSENSITIVE2 (KAI2), functions as a KAR receptor and likely recognizes an uncharacterized, endogenous signal ('KL'). Previous phylogenetic analyses have suggested that the KAI2 lineage is ancestral in land plants, and that canonical D14-type SL receptors only arose in seed plants; this is paradoxical, however, as non-vascular plants synthesize and respond to SLs. Results: We have used a combination of phylogenetic and structural approaches to re-assess the evolution of the D14/KAI2 family in land plants. We analysed 339 members of the D14/KAI2 family from land plants and charophyte algae. Our phylogenetic analyses show that the divergence between the eu-KAI2 lineage and the DDK (D14/DLK2/ KAI2) lineage that includes D14 occurred very early in land plant evolution. We show that eu-KAI2 proteins are highly conserved, and have unique features not found in DDK proteins. Conversely, we show that DDK proteins show considerable sequence and structural variation to each other, and lack clearly definable characteristics. We use homology modelling to show that the earliest members of the DDK lineage structurally resemble KAI2 and that SL receptors in non-seed plants likely do not have D14-like structure. We also show that certain groups of DDK proteins lack the otherwise conserved MORE AXILLARY GROWTH2 (MAX2) interface, and may thus function independently of MAX2, which we show is highly conserved throughout land plant evolution. Conclusions: Our results suggest that D14-like structure is not required for SL perception, and that SL perception has relatively relaxed structural requirements compared to KAI2-mediated signalling. We suggest that SL perception gradually evolved by neo-functionalization within the DDK lineage, and that the transition from KAI2-like to D14-like protein may have been driven by interactions with protein partners, rather than being required for SL perception per se. [ABSTRACT FROM AUTHOR]

Strigolactones are a structurally diverse class of plant hormones that control many aspects of shoot and root growth. Strigolactones are also exuded by plants into the rhizosphere, where they promote symbiotic interactions with arbuscular mycorrhizal fungi and germination of root parasitic plants in the Orobanchaceae family. Therefore, understanding how strigolactones are made, transported, and perceived may lead to agricultural innovations as well as a deeper knowledge of how plants function. Substantial progress has been made in these areas over the past decade. In this review, we focus on the molecular mechanisms, core developmental roles, and evolutionary history of strigolactone signaling. We also propose potential translational applications of strigolactone research to agriculture. [ABSTRACT FROM AUTHOR]

Strigolactones (SLs) are hormonal signals that regulate multiple aspects of shoot architecture, including shoot branching. Like many plant hormonal signaling systems, SLs act by promoting ubiquitination of target proteins and their subsequent proteasome-mediated degradation. Recently, SMXL6, SMXL7, and SMXL8, members of the SMAX1-LIKE (SMXL) family of chaperonin-like proteins, have been identified as proteolytic targets of SL signaling in Arabidopsis thaliana. However, the mechanisms by which these proteins regulate downstream events remain largely unclear. Here, we show that SMXL7 functions in the nucleus, as does the SL receptor, DWARF14 (D14). We show that nucleus-localized D14 can physically interact with both SMXL7 and the MAX2 F-box protein in a SL-dependent manner and that disruption of specific conserved domains in SMXL7 affects its localization, SL-induced degradation, and activity. By expressing and overexpressing these SMXL7 protein variants, we show that shoot tissues are broadly sensitive to SMXL7 activity, but degradation normally buffers the effect of increasing SMXL7 expression. SMXL7 contains a well-conserved EAR (ETHYLENE-RESPONSE FACTOR Amphiphilic Repression) motif, which contributes to, but is not essential for, SMXL7 functionality. Intriguingly, different developmental processes show differential sensitivity to the loss of the EAR motif, raising the possibility that there may be several distinct mechanisms at play downstream of SMXL7. [ABSTRACT FROM AUTHOR]

The plant hormones strigolactones and smoke-derived karrikins are butenolide signals that control distinct aspects of plant development. Perception of both molecules in Arabidopsis thaliana requires the F-box protein MORE AXILLARY GROWTH2 (MAX2). Recent studies suggest that the homologous SUPPRESSOR OF MAX2 1 (SMAX1) in Arabidopsis and DWARF53 (D53) in rice (Oryza sativa) are downstream targets of MAX2. Through an extensive analysis of loss-of-function mutants, we demonstrate that the Arabidopsis SMAX1-LIKE genes SMXL6 , SMXL7 , and SMXL8 are co-orthologs of rice D53 that promote shoot branching. SMXL7 is degraded rapidly after treatment with the synthetic strigolactone mixture rac -GR24. Like D53, SMXL7 degradation is MAX2- and D14-dependent and can be prevented by deletion of a putative P-loop. Loss of SMXL6,7,8 suppresses several other strigolactone-related phenotypes in max2 , including increased auxin transport and PIN1 accumulation, and increased lateral root density. Although only SMAX1 regulates germination and hypocotyl elongation, SMAX1 and SMXL6,7,8 have complementary roles in the control of leaf morphology. Our data indicate that SMAX1 and SMXL6,7,8 repress karrikin and strigolactone signaling, respectively, and suggest that all MAX2-dependent growth effects are mediated by degradation of SMAX1/SMXL proteins. We propose that functional diversification within the SMXL family enabled responses to different butenolide signals through a shared regulatory mechanism. [ABSTRACT FROM AUTHOR]

There has been a dramatic recent increase in the understanding of the mechanisms by which plants detect their neighbors, 1 including by touch, 2 reflected light, 3 volatile organic chemicals, and root exudates. 4,5 The importance of root exudates remains ill-defined because of confounding experimental variables 6,7 and difficulties disentangling neighbor detection in shoot and roots. 8–10 There is evidence that root exudates allow distinction between kin and non-kin neighbors, 11–13 but identification of specific exudates that function in neighbor detection and/or kin recognition remain elusive. 1 Strigolactones (SLs), which are exuded into the soil in significant quantities in flowering plants to promote recruitment of arbuscular mycorrhizal fungi (AMF), 14 seem intuitive candidates to act as plant-plant signals, since they also act as hormones in plants, 15–17 with dramatic effects on shoot growth 18,19 and milder effects on root development. 20 Here, using pea, we test whether SLs act as either cues or signals for neighbor detection. We show that peas detect neighbors early in the life cycle through their root systems, resulting in strong changes in shoot biomass and branching, and that this requires SL biosynthesis. We demonstrate that uptake and detection of SLs exuded by neighboring plants are needed for this early neighbor detection, and that plants that cannot exude SLs are outcompeted by neighboring plants and fail to adjust growth to their soil volume. We conclude that plants both exude SLs as signals to modulate neighbor growth and detect environmental SLs as a cue for neighbor presence; collectively, this allows plants to proactively adjust their shoot growth according to neighbor density. • Pea plants detect neighbors through their root systems early in life • Strigolactone exudation by neighbors is needed for this response • Strigolactone exudation mutants are outcompeted by their neighbors • Environmental strigolactones thus act as cue for neighbor presence Wheeldon et al. show that pea plants change shoot growth in response to strigolactones exuded by neighboring plant roots. They show that plants that cannot exude strigolactones are outcompeted by neighbors and conclude that environmental strigolactones therefore act as a cue for neighbor presence in plants. [ABSTRACT FROM AUTHOR]

Plants have evolved elaborate mechanisms to detect neighboring plants, which typically involve the perception of "cues" inadvertently produced by the neighbor. 1 Strigolactones are hormonal signaling molecules 2,3 that are also exuded into the rhizosphere by most flowering plant species to promote arbuscular mycorrhizal symbioses. 4 Since flowering plants have an endogenous perception system for strigolactones, 5 strigolactones are obvious candidates to act as a cue for neighbor presence, but have not been shown to act as such. To test this hypothesis in rice plants, we quantified two major strigolactones of rice plants, orobanchol and 4-deoxyorobanchol, in root exudates by using LC-MS/MS (MRM) and examined feedback regulation of strigolactone biosynthesis and changes in shoot branching phenotypes in rice plants grown at different densities in hydroponics and soil culture. We show that the presence of neighboring plants, or greater root volume, results in rapidly induced changes in strigolactone biosynthesis, sensitivity, and exudation and the subsequent longer-term changes in shoot architecture. These changes require intact strigolactone biosynthesis in neighboring plants and intact strigolactone signaling in focal plants. These results suggest that strigolactone biosynthesis and exudation in rice plants are driven by supra-organismal environmental strigolactone levels. Strigolactones thus act as a cue for neighbor presence in rice plants, but also seem to act as a more general root density-sensing mechanism in flowering plants that integrates soil volume and neighbor density and allows plants to adapt to the limitations of the rhizosphere. • Crowding reduces biosynthesis and exudation of strigolactones • This density-dependent response requires strigolactone perception • Strigolactone biosynthesis and exudation respond to environmental strigolactones • Crowding in soil reduces strigolactone exudation and shoot branching in rice plants Strigolactones (SLs) trigger parasitism and symbiosis in the rhizosphere and function as a hormone in plants. Yoneyama et al. demonstrate that rice plants sense SLs in the rhizosphere and regulate their SL biosynthesis and exudation, suggesting that SLs act as a cue for a plant-plant communication in rice plants. [ABSTRACT FROM AUTHOR]