Cytokinins (CKs) and glucose (GLC) control a number of common responses in plants. We hypothesize that there may be an extensive overlap between CK-and GLC-signalling pathways. Microarray along with physiological analysis has been performed to find out the interdependence/overlap between CK and GLC signal transduction pathways in Arabidopsis seedlings. GLC could transcriptionally affect 76% of CK-regulated genes at whole genome level, 89% of which are agonistically regulated. GLC may also affect CK-regulated gene expression via non-transcriptional pathways. GLC can regulate several genes involved in CK metabolism and signalling. A number of gene families involved in development and stress are commonly regulated by CK and GLC. Physiologically, both GLC and CK could regulate hypocotyl length in dark. GLC and CK signalling may integrate at the level of type A Arabidopsis response regulators (ARRs) in controlling hypocotyl length. Both GLC and CK signalling cannot alter hypocotyl length in dark in auxin-signalling mutants AUXIN RESPONSE2/INDOLE-3-ACETIC ACID7 (AXR2/IAA7) and AXR3/IAA17 suggesting that they may involve auxinsignalling component as a nodal point. Here, we demonstrate that there is an extensive overlap between CK-and GLCregulated gene expression and physiological responses.

(A.M.J.) Optimal root architecture is established by multiple intrinsic (e.g. hormones) and extrinsic (e.g. gravity and touch) signals and is established, in part, by directed root growth. We show that asymmetrical exposure of cytokinin (CK) at the root tip in Arabidopsis (Arabidopsis thaliana) promotes cell elongation that is potentiated by glucose in a hexokinase-influenced, G proteinindependent manner. This mode of CK signaling requires the CK receptor, ARABIDOPSIS HISTIDINE KINASE4 and, at a minimum, its cognate type B ARABIDOPSIS RESPONSE REGULATORS ARR1, ARR10, and ARR11 for full responsiveness, while type A response regulators act redundantly to attenuate this CK response. Ethylene signaling through the ethylene receptor ETHYLENE RESISTANT1 and its downstream signaling element ETHYLENE INSENSITIVE2 are required for CKinduced root cell elongation. Negative and positive feedback loops are reinforced by CK regulation of the expression of the genes encoding these elements in both the CK and ethylene signaling pathways. Auxin transport facilitated by PIN-FORMED2 as well as auxin signaling through control of the steady-state level of transcriptional repressors INDOLE-3-ACETIC ACID7 (IAA7), IAA14, and IAA17 via TRANSPORT INHIBITOR RESPONSE1/AUXIN SIGNALING F-BOX PROTEIN are involved in CK-induced root cell elongation. This action lies downstream of ethylene and CK induction. Intrinsic signaling in this response operates independently of the extrinsic signal touch, although actin filament organization, which is important in the touch response, may be important for this response, since latrunculin B can induce similar growth. This root growth response may have adaptive significance, since CK responsiveness is inversely related to root coiling and waving, two root behaviors known to be important for fitness.

Xyloglucan is the major hemicellulose of dicotyledon primary cell walls, affecting the load-bearing framework with the participation of xyloglucan endo-transglycosylase/hydrolases (XTHs). We used loss-and gain-of function approaches to study functions of XTH4 and XTH9 abundantly expressed in cambial regions during secondary growth of Arabidopsis (Arabidopsis thaliana). In secondarily thickened hypocotyls, these enzymes had positive effects on vessel element expansion and fiber intrusive growth. They also stimulated secondary wall thickening but reduced secondary xylem production. Cell wall analyses of inflorescence stems revealed changes in lignin, cellulose, and matrix sugar composition indicating an overall increase in secondary versus primary walls in mutants, indicative of higher xylem production compared with the wild type (since secondary walls were thinner). Intriguingly, the number of secondary cell wall layers compared with the wild type was increased in xth9 and reduced in xth4, whereas the double mutant xth4x9 displayed an intermediate number of layers. These changes correlated with specific Raman signals from the walls, indicating changes in lignin and cellulose. Secondary walls were affected also in the interfascicular fibers, where neither XTH4 nor XTH9 was expressed, indicating that these effects were indirect. Transcripts involved in secondary wall biosynthesis and cell wall integrity sensing, including THESEUS1 and WALL ASSOCIATED KINASE2, were highly induced in the mutants, indicating that deficiency in XTH4 and XTH9 triggers cell wall integrity signaling, which, we propose, stimulates xylem cell production and modulates secondary wall thickening. Prominent effects of XTH4 and XTH9 on secondary xylem support the hypothesis that altered xyloglucan affects wood properties both directly and via cell wall integrity sensing. cell wall composition, mutant growth, and gene expression and wrote the article; A.B. analyzed hypocotyl phenotypes and xylem cell morphology in mutants, discovered secondary wall phenotypes, and carried out immunolocalization; N.N. isolated and purified mutants and created OE plants; M.D.-M. and M.M. analyzed cell walls by in situ Raman and in situ FTIR; S.E. created GUS reporter lines; V.K. analyzed Populus XTH genes and their expression in Asp-Wood; L.G. and S.M.-M. carried out saccharification analyses; A.G. interpreted FTIR and Raman data and helped with spectroscopic analyses; J.B. analyzed XET activity; B.S. and E.J.M. conceived and coordinated the project; E.J.M. finalized the article with contribution from all authors.[OPEN]Articles can be viewed without a subscription. www.plantphysiol.org/cgi/

(E.J.M.). Tyloses are ingrowths of parenchyma cells into the lumen of embolized xylem vessels, thereby protecting the remaining xylem from pathogens. They are found in heartwood, sapwood, and in abscission zones and can be induced by various stresses, but their molecular triggers are unknown. Here, we report that down-regulation of PECTIN METHYLESTERASE1 (PtxtPME1) in aspen (Populus tremula 3 tremuloides) triggers the formation of tyloses and activation of oxidative stress. We tested whether any of the oxidative stress-related hormones could induce tyloses in intact plantlets grown in sterile culture. Jasmonates, including jasmonic acid (JA) and methyl jasmonate, induced the formation of tyloses, whereas treatments with salicylic acid (SA) and 1-aminocyclopropane-1-carboxylic acid (ACC) were ineffective. SA abolished the induction of tyloses by JA, whereas ACC was synergistic with JA. The ability of ACC to stimulate tyloses formation when combined with JA depended on ethylene (ET) signaling, as shown by a decrease in the response in ET-insensitive plants. Measurements of internal ACC and JA concentrations in wild-type and ET-insensitive plants treated simultaneously with these two compounds indicated that ACC and JA regulate each other's concentration in an ET-dependent manner. The findings indicate that jasmonates acting synergistically with ethylene are the key molecular triggers of tyloses.