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Cytokinin hormones are important regulators of development and environmental responses of plants that execute their action via the molecular machinery of signal perception and transduction. The limiting step of the whole process is the availability of the hormone in suitable concentrations in the right place and at the right time to interact with the specific receptor. Hence, the hormone concentrations in individual tissues, cells, and organelles must be properly maintained by biosynthetic and metabolic enzymes. Although there are merely two active cytokinins, isopentenyladenine and its hydroxylated derivative zeatin, a variety of conjugates they may form and the number of enzymes/isozymes with varying substrate specificity involved in their biosynthesis and conversion gives the plant a variety of tools for fine tuning of the hormone level. Recent genome-wide studies revealed the existence of the respective coding genes and gene families in plants and in some bacteria. This review summarizes present knowledge on the enzymes that synthesize cytokinins, form cytokinin conjugates, and carry out irreversible elimination of the hormones, including their phylogenetic analysis and possible variations in different organisms.

Summary The cytokinins, which are N6‐substituted adenine derivatives, control key aspects of crop productivity. Cytokinin levels are controlled via biosynthesis by isopentenyl transferase (IPT), destruction by cytokinin oxidase/dehydrogenase (CKX), and inactivation via glucosylation by cytokinin glucosyl transferases (CGTs). While both yield components and tolerance to drought and related abiotic stressors have been positively addressed via manipulation of IPT and/or CKX expression, much less attention has been paid to the CGTs. As naming of the CGTs has been unclear, we suggest COGT, CNGT, CONGT and CNOGT to describe the O‐, N‐ and dual function CGTs. As specific CGT mutants of both rice and arabidopsis showed impacts on yield components, we interrogated the wheat genome database, IWGSC RefSeq v1.0 & v2.0, to investigate wheat CGTs. Besides providing unambiguous names for the 53 wheat CGTs, we show their expression patterns in 70 developmental tissues and their response characteristics to various stress conditions by reviewing more than 1000 RNA‐seq data sets. These revealed various patterns of responses and showed expression generally being more limited in reproductive tissues than in vegetative tissues. Multiple cis‐regulatory elements are present in the 3 kb upstream of the start codons of the 53 CGTs. Elements associated with abscisic acid, light and methyl jasmonate are particularly over‐represented, indicative of the responsiveness of CGTs to the environment. These data sets indicate that CGTs have potential value for wheat improvement and that these could be targeted in TILLING or gene editing wheat breeding programmes.

Increasing shoot branch numbers of Chrysanthemum morifolium Ramat cv. Hangbaiju (Hangbaiju) is crucial for producing high flower yields. Pot experiments were designed to evaluate the effects of foliar application of 6-benzylaminopurine (6-BA) combined with prohexadione-Ca (Pro-Ca) on the yield and quality of Hangbaiju flowers. Foliar application of 6-BA combined with Pro-Ca typically increased leaf chlorophyll content and decreased leaf soluble sugar and soluble protein contents throughout the floral organ growth phase. At the bud formation phenophase stage (August), the contents of gibberellin (GA), indole-3-acetic acid (IAA), and zeatin (ZA) decreased (except for GA content in the 10 mg L[sup.−1] 6-BA combined with 100 mg L[sup.−1] Pro-Ca treatment), but abscisic acid (ABA) content increased. The yield of Hangbaiju flowers was found higher in plants treated with foliar application of 6-BA combined with Pro-Ca. Higher yields were found in the 6-BA[sub.5] + Ca[sub.100] and 6-BA[sub.10] + Ca[sub.100] treatments than in the 6-BA[sub.5] + Ca[sub.50] and 6-BA[sub.10] + Ca[sub.50] treatments, and the highest yield was observed in the 6-Ba[sub.5] + Ca[sub.100] treatment both in 2019 and 2020. Nutritional indices such as soluble sugar, soluble protein, total amino acid, and water extract increased, and medicinal indices such as flavonoid, total phenolics, chlorogenic acid, cynaroside, and 3,5-dicaffeoyl quinic acid slightly decreased after foliar application of 6-BA combined with Pro-Ca; however, they were all higher than the standards recorded in the Chinese Pharmacopeia. Overall, foliar application of 6-BA combined with Pro-Ca could increase the yield and nutritional quality of Hangbaiju flowers, enhancing its tea quality.

Cytokinins are plant hormones, derivatives of adenine with a side chain at the N6-position. They are involved in many physiological processes. While the metabolism of trans-zeatin and isopentenyladenine, which are considered to be highly active cytokinins, has been extensively studied, there are others with less obvious functions, such as cis-zeatin, dihydrozeatin, and aromatic cytokinins, which have been comparatively neglected. To help explain this duality, we present a novel hypothesis metaphorically comparing various cytokinin forms, enzymes of CK metabolism, and their signalling and transporter functions to the comics superheroes Hulk and Deadpool. Hulk is a powerful but short-lived creation, whilst Deadpool presents a more subtle and enduring force. With this dual framework in mind, this review compares different cytokinin metabolites, and their biosynthesis, translocation, and sensing to illustrate the different mechanisms behind the two CK strategies. This is put together and applied to a plant developmental scale and, beyond plants, to interactions with organisms of other kingdoms, to highlight where future study can benefit the understanding of plant fitness and productivity.

• The diversity of cytokinin (CK) metabolites suggests their interconversions are the predominant regulatory mechanism of CK action. Nevertheless, little is known about their directionality and kinetics in planta. • CK metabolite levels were measured in 2-wk-old Arabidopsis thaliana plants at several time points up to 100 min following exogenous application of selected CKs. The data were then evaluated qualitatively and by mathematical modeling. • Apart from elevated levels of trans-zeatin (tZ) metabolites upon application of N⁶-(Δ²-isopentenyl)adenine (iP), we observed no conversions between the individual CK-types – iP, tZ, dihydrozeatin (DHZ) and cis-zeatin (cZ). In particular, there was no sign of isomerization between tZ and cZ families. Also, no increase of DHZ-type CKs was observed after application of tZ, suggesting low baseline activity of zeatin reductase. Among N-glucosides, those of iP were not converted back to iP while tZ N-glucosides were cleaved to tZ bases, thus affecting the whole metabolic spectrum. • We present the first large-scale study of short-term CK metabolism kinetics and show that tZ N7- and N9-glucosides are metabolized in vivo. We thus refute the generally accepted hypothesis that N-glucosylation irreversibly inactivates CKs. The subsequently constructed mathematical model provides estimates of the metabolic conversion rates.

The metabolism of cytokinins (CKs) and auxins in vascular plants is relatively well understood, but data concerning their metabolic pathways in non-vascular plants are still rather rare. With the aim of filling this gap, 20 representatives of taxonomically major lineages of cyanobacteria and algae from Cyanophyceae, Xanthophyceae, Eustigmatophyceae, Porphyridiophyceae, Chlorophyceae, Ulvophyceae, Trebouxiophyceae, Zygnematophyceae and Klebsormidiophyceae were analysed for endogenous profiles of CKs and auxins and some of them were used for studies of the metabolic fate of exogenously applied radiolabelled CK, [ H]trans-zeatin (transZ) and auxin ([ H]indole-3-acetic acid (IAA)), and the dynamics of endogenous CK and auxin pools during algal growth and cell division. Quantification of phytohormone levels was performed by high-performance or ultrahigh-performance liquid chromatography-electrospray tandem mass spectrometry (HPLC-MS/MS, UHPLC-MS/MS). The dynamics of exogenously applied [ H]transZ and [ H]IAA in cell cultures were monitored by HPLC with on-line radioactivity detection. The comprehensive screen of selected cyanobacteria and algae for endogenous CKs revealed a predominance of bioactive and phosphate CK forms while O- and N-glucosides evidently did not contribute greatly to the total CK pool. The abundance of cis-zeatin-type CKs and occurrence of CK 2-methylthio derivatives pointed to the tRNA pathway as a substantial source of CKs. The importance of the tRNA biosynthetic pathway was proved by the detection of tRNA-bound CKs during the course of Scenedesmus obliquus growth. Among auxins, free IAA and its oxidation catabolite 2-oxindole-3-acetic acid represented the prevailing endogenous forms. After treatment with [ H]IAA, IAA-aspartate and indole-3-acetyl-1-glucosyl ester were detected as major auxin metabolites. Moreover, different dynamics of endogenous CKs and auxin profiles during S. obliquus culture clearly demonstrated diverse roles of both phytohormones in algal growth and cell division. Our data suggest the existence and functioning of a complex network of metabolic pathways and activity control of CKs and auxins in cyanobacteria and algae that apparently differ from those in vascular plants.

Cytokinin signaling plays an important role in plant growth and development, and therefore its molecular characteristics are under extensive study. One characteristic is the subcellular localization of cytokinin signal initiation. This localization determines both the pathway for hormone delivery to the receptor, as well as molecular aspects of signal transfer to the primary cellular targets. Subcellular sites for the onset of cytokinin signaling are still uncertain and experimental data are in part controversial. A few years ago, cytokinin receptors were shown to be localized predominantly in the membrane of the endoplasmic reticulum (ER) and to possess some features, such as their pH activity profile, typical for intracellular proteins. Very recently, new data corroborating the functionality of ER-located cytokinin receptors were reported. However, other work argued for cytokinin perception to occur at the plasma membrane (PM). Here, we discuss in detail these partially conflicting data and present an integrative model for cytokinin perception and signaling. In our opinion, the prevailing evidence argues for the ER being the predominant site of cytokinin signal perception but also that signal initiation at the PM might be relevant in some circumstances as well. The roles of these pathways in long-distance, paracrine and autocrine cytokinin signaling are discussed.

Cytokinins are important signals that participate in different plant processes, and are well known for their strong influence in plant development. With the years, knowledge has been built about their effects, chemical nature, metabolism, and signaling mechanisms. However, one aspect about cytokinins that has been lagging behind is cytokinin transport. Recent reports are providing more information about how cytokinins are transported and how their transport is connected to their effects in development. This review provides a general overview of what is known about cytokinin transport, with a focus on the latest reports.

Strigolactones (SLs), a group of terpenoid lactones derived from carotenoids, are plant hormones that control numerous aspects of plant development. Although the framework of SL signaling that the repressor DWARF 53 (D53) could be SL-dependently degraded via the SL receptor D14 and F-box protein D3 has been established, the downstream response genes to SLs remain to be elucidated. Here we show that the cytokinin (CK) content is dramatically increased in shoot bases of the rice SL signaling mutant d53. By examining transcript levels of all the CK metabolism-related genes after treatment with SL analog GR24, we identified CYTOKININ OXIDASE/DEHYDROGENASE 9 (OsCKX9) as a primary response gene significantly up-regulated within 1 h of treatment in the wild type but not in d53. We also found that OsCKX9 functions as a cytosolic and nuclear dual-localized CK catabolic enzyme, and that the overexpression of OsCKX9 suppresses the browning of d53 calli. Both the CRISPR/Cas9-generated OsCKX9 mutants and OsCKX9-overexpressing transgenic plants showed significant increases in tiller number and decreases in plant height and panicle size, suggesting that the homeostasis of OsCKX9 plays a critical role in regulating rice shoot architecture. Moreover, we identified the CK-inducible rice type-A response regulator OsRR5 as the secondary SL-responsive gene, whose expression is significantly repressed after 4 h of GR24 treatment in the wild type but not in osckx9. These findings reveal a comprehensive plant hormone cross-talk in which SL can induce the expression of OsCKX9 to down-regulate CK content, which in turn triggers the response of downstream genes.