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Cytokinin (CK) has long been implicated as a promoter of bud outgrowth in plants, but exactly how this is achieved in coordination with other plant hormones is unclear. The recent discovery of strigolactones (SLs) as the long-sought branch-inhibiting hormone allowed us to test how CK and SL coordinately regulate bud outgrowth in pea (Pisum sativum). We found that SL-deficient plants are more sensitive to stimulation of bud growth by low concentrations of locally applied CK than wildtype plants. Furthermore, in contrast with SL mutant plants, buds of wild-type plants are almost completely resistant to stimulation by CK supplied to the vasculature. Regardless of whether the exogenous hormones were supplied locally or to the xylem stream, SL and CK acted antagonistically on bud outgrowth. These data suggest that SLs do not affect the delivery of CK to axillary buds and vice versa. Rather, these data combined with dose-response experiments suggest that SLs and CK can act directly in buds to control their outgrowth. These hormones may converge at a common point in the bud outgrowth regulatory pathway. The expression of pea BRANCHED1, a TCP transcription factor expressed strongly in buds and thought to act downstream of SLs in shoot branching, is regulated by CK and SL without a requirement for protein synthesis and in a manner that correlates with observed bud growth responses.

Strigolactones (SLs) are key hormonal regulators of flowering plant development and are widely distributed amongst streptophytes. In Arabidopsis, SLs signal via the F-box protein MORE AXILLARY GROWTH2 (MAX2), affecting multiple aspects of development including shoot branching, root architecture and drought tolerance. Previous characterization of a Physcomitrella patens moss mutant with defective SL synthesis supports an ancient role for SLs in land plants, but the origin and evolution of signalling pathway components are unknown. Here we investigate the function of a moss homologue of MAX2, PpMAX2, and characterize its role in SL signalling pathway evolution by genetic analysis. We report that the moss Ppmax2 mutant shows very distinct phenotypes from the moss SL-deficient mutant. In addition, the Ppmax2 mutant remains sensitive to SLs, showing a clear transcriptional SL response in dark conditions, and the response to red light is also altered. These data suggest divergent evolutionary trajectories for SL signalling pathway evolution in mosses and vascular plants. In P.patens, the primary roles for MAX2 are in photomorphogenesis and moss early development rather than in SL response, which may require other, as yet unidentified, factors.

Strigolactone (SL) mutants in diverse species show reduced stature in addition to their extensive branching. Here, we show that this dwarfism in pea (Pisum sativum) is not attributable to the strong branching of the mutants. The continuous supply of the synthetic SL GR24 via the root system using hydroponics can restore internode length of the SL-deficient rms1 mutant but not of the SL-response rms4 mutant, indicating that SLs stimulate internode elongation via RMS4. Cytological analysis of internode epidermal cells indicates that SLs control cell number but not cell length, suggesting that SL may affect stem elongation by stimulating cell division. Consequently, SLs can repress (in axillary buds) or promote (in the stem) cell division in a tissue-dependent manner. Because gibberellins (GAs) increase internode length by affecting both cell division and cell length, we tested if SLs stimulate internode elongation by affecting GA metabolism or signaling. Genetic analyses using SL-deficient and GA-deficient or DELLA-deficient double mutants, together with molecular and physiological approaches, suggest that SLs act independently from GAs to stimulate internode elongation.

The function of PsBRCl, the pea (Pisum sativum) homolog of the maize (Zea mays) TEOSINTE BRANCHED1 and the Arabidopsis (Arabidopsis thaliana) BRANCHED1 (AtBRC1) genes, was investigated. The pea Psbrc1 mutant displays an increased shoot-branching phenotype, is able to synthesize strigolactone (SL), and does not respond to SL application. The level of pleiotropy of the SL-deficient ramosus1 (rms1) mutant is higher than in the Psbrcl mutant, rms1 exhibiting a relatively dwarf phenotype and more extensive branching at upper nodes. The PsBRCl gene is mostly expressed in the axillary bud and is transcriptionally up-regulated by direct application of the synthetic SL GR24 and down-regulated by the cytokinin (CK) 6-benzylaminopurine. The results suggest that PsBRCl may have a role in integrating SL and CK signals and that SLs act directly within the bud to regulate its outgrowth. However, the Psbrcl mutant responds to 6-benzylaminopurine application and decapitation by increasing axillary bud length, implicating a PsBRCl -independent component of the CK response in sustained bud growth. In contrast to other SL-related mutants, the Psbrcl mutation does not cause a decrease in the CK zeatin riboside in the xylem sap or a strong increase in RMS1 transcript levels, suggesting that the RMS2-dependent feedback is not activated in this mutant. Surprisingly, the double rmsl Psbrcl mutant displays a strong increase in numbers of branches at cotyledonary nodes, whereas branching at upper nodes is not significantly higher than the branching in rmsl. This phenotype indicates a localized regulation of branching at these nodes specific to pea.

Initially known for their role in the rhizosphere in stimulating the seed germination of parasitic weeds such as the Striga and Orobanche species, and later as host recognition signals for arbuscular mycorrhizal fungi, strigolactones (SLs) were recently rediscovered as a new class of plant hormones involved in the control of shoot branching in plants. Herein, we report the synthesis of new SL analogs and, to our knowledge, the first study of SL structure-activity relationships for their hormonal activity in garden pea (Pisum sativum). Comparisons with their action for the germination of broomrape (Phelipanche ramosa) are also presented. The pea rms1 SL-deficient mutant was used in a SL bioassay based on axillary bud length after direct SL application on the bud. This assay was compared with an assay where SLs were fed via the roots using hydroponics and with a molecular assay in which transcript levels of BRANCHED1, the pea homolog of the maize TEOSINTE BRANCHED1 gene were quantified in axillary buds only 6 h after application of SLs. We have demonstrated that the presence of a Michael acceptor and a methylbutenolide or dimethylbutenolide motif in the same molecule is essential. It was established that the more active analog 23 with a dimethylbutenolide as the D-ring could be used to control the plant architecture without strongly favoring the germination of P. ramosa seeds. Bold numerals refer to numbers of compounds.

Uncovering the basis of small molecule hormone receptors evolution is paramount to a complete understanding of how protein structure drives function. In plants, hormone receptors for strigolactones are well suited to evolutionary inquires because closely related homologs have different ligand preferences. More importantly, because of facile plant transgenic systems, receptors can be swapped and quickly assessed functionally in vivo. Here, we show only three mutations are required to switch the non-strigolactone receptor, KAI2, into a strigolactone receptor. This modified receptor still perceives KAI2 ligands and does not require receptor hydrolysis for activity. Structural and molecular dynamic modeling suggest receptor pocket flexibility is important for ligand specificity and downstream signaling partner affinity. These findings indicate a few keystone mutations link strigolactone signaling to germination, which explains how parasitic plants that devastate African agriculture evolved SL receptors to sense the presence of a host plant.

The timing of leaf emergence at the shoot apical meristem, or plastochron, is highly regulated in plants. Among the genes known to regulate the plastochron in Arabidopsis (Arabidopsis thaliana), KLUH (KLU), orthologous to the rice (Oryza sativa) PLASTOCHRON1, encodes the cytochrome P450 CYP78A5, and is thought to act through generation of a still unknown mobile signal. As klu mutants display not only a short plastochron but also a branching phenotype reminiscent of strigolactone (SL) mutants, we investigated whether KLU/CYP78A5 is involved in SL biosynthesis. We combined a genetic approach, a parasitic plant seed germination bioassay to test klu root exudates, and analysis of transcript abundances of SL-biosynthesis genes in the Arabidopsis klu mutants. We demonstrate that KLU is not involved in the SL-biosynthesis pathway. Moreover, this work allowed us to uncover a new role for SL during Arabidopsis development in modulating plastochron via a KLU-dependent pathway. Globally our data reveal that KLU is required for plastochron-specific SL responses, a first indication of crosstalk between SL and the KLU-derived signal.

Although the benefits of physical exercise to preserve bone quality are now widely recognized, the intimate mechanisms leading to the underlying cell responses still require further investigations. Interval training running, for instance, appears as a generator of impacts on the skeleton, and particularly on the progenitor cells located in the bone marrow. Therefore, if this kind of stimulus initiates bone cell proliferation and differentiation, the activation of a devoted signaling pathway by mechano-transduction seems likely. This study aimed at investigating the effects of an interval running program on the appearance of the zinc finger protein FHL2 in bone cells and their anatomical location. Twelve 5-week-old male Wistar rats were randomly allocated to one of the following groups (n = 6 per group): sedentary control (SED) or high-intensity interval running (EX, 8 consecutive weeks). FHL2 identification in bone cells was performed by immuno-histochemistry on serial sections of radii. We hypothesized that impacts generated by running could activate, in vivo, a specific signaling pathway, through an integrin-mediated mechano-transductive process, leading to the synthesis of FHL2 in bone marrow cells. Our data demonstrated the systematic appearance of FHL2 (% labeled cells: 7.5%, p < 0.001) in bone marrow obtained from EX rats, whereas no FHL2 was revealed in SED rats. These results suggest that the mechanical impacts generated during high-intensity interval running activate a signaling pathway involving nuclear FHL2, such as that also observed with dexamethasone administration. Consequently, interval running could be proposed as a non-pharmacological strategy to contribute to bone marrow cell osteogenic differentiation.

The natural growth regulator zaxinone increases the levels of the phytohormones strigolactone (SL) and abscisic acid in Arabidopsis ( Arabidopsis thaliana ) via unknown mechanisms. Here we demonstrate that part of the effect of zaxinone in Arabidopsis depend on the SL receptor DWARF14 ( At D14) and the F-Box protein MORE AXILLARY BRANCHING2 ( At MAX2) that mediate the signaling of SLs and karrikins. Binding assays and co-crystallization reveal zaxinone as an additional ligand of At D14 and an SL antagonist that interrupts the interaction of At D14 with At MAX2. Zaxinone also binds to the karrikin receptor KARRIKIN INSENSITIVE2 ( At KAI2). These findings unveil a perception mechanism for zaxinone in Arabidopsis and demonstrate the capability of At D14 to bind signaling molecules, other than strigolactones and mediate their transduction. Zaxinone is a novel ligand of the strigolactone (SL) receptor DWARF14 (D14) in Arabidopsis. It competitively binds the active site of D14 and acts as a long-lasting, non-hydrolysable antagonist of SL by blocking the interaction between D14 and MAX2.

Purpose To determine whether isokinetic muscle recovery following ACLR using a hamstring tendon (HT) would be equivalent (non-inferior) in knees that had high-grade pivot-shift and adjuvant modified Lemaire procedure versus knees that had minimal pivot-shift and no adjuvant modified Lemaire procedure. Methods We evaluated 96 consecutive patients that underwent primary ACLR. Nine were excluded because of contralateral knee injury, and of the remaining 87, ACLR was performed stand-alone in 52 (Reference group), and with a Lemaire procedure in 35 (Lemaire group) who had high-grade pivot-shift, age < 18, or genu recurvatum > 20°. At 6 months, isokinetic tests were performed at 240°/s and 90°/s to calculate strength deficits of hamstrings (H) and quadriceps (Q). At 8 months, patients were evaluated using IKDC, Lysholm, and Tegner scores. Results Compared to the Reference group, the Lemaire group were younger (23.0 ± 2.5 vs 34.2 ± 10.5, p  = 0.021) with a greater proportion of males (80% vs 56%, p  < 0.001). The Lemaire group had no complications, but the Reference group had one graft failure and one cyclops syndrome. Strength deficits at 240°/s and at 90°/s were similar in both groups, but mixed H/Q ratios were lower for the Lemaire group (1.02 ± 0.19 vs 1.14 ± 0.24, p  = 0.011). IKDC and Lysholm scores were similar in both groups, but Tegner scores were higher in the Lemaire group (median, 6.5 vs 6.0, p  = 0.024). Conclusions ACLR with a modified Lemaire procedure for knees with rotational instability grants equivalent isokinetic muscle recovery as stand-alone ACLR in knees with no rotational instability. For ACL-deficient knees with high-grade pivot-shift, a Lemaire procedure restores rotational stability without compromising isokinetic muscle recovery. Study design Level III, comparative study.