Optimizing root system architecture can overcome yield limitations in crop plants caused by water or nutrient shortages. Classic breeding approaches are difficult because the trait is governed by many genes and is difficult to score. We generated transgenic Arabidopsis thaliana and tobacco (Nicotiana tabacum) plants with enhanced root-specific degradation of the hormone cytokinin, a negative regulator of root growth. These transgenic plants form a larger root system, whereas growth and development of the shoot are similar. Elongation of the primary root, root branching, and root biomass formation were increased by up to 60% in transgenic lines, increasing the root-to-shoot ratio. We thus demonstrated that a single dominant gene could regulate a complex trait, root growth. Moreover, we showed that cytokinin regulates root growth in a largely organ-autonomous fashion that is consistent with its dual role as a hormone with both paracrine and long-distance activities. Transgenic plants had a higher survival rate after severe drought treatment. The accumulation of several elements, including S, P, Mn, Mg, Zn, as well as Cd from a contaminated soil, was significantly increased in shoots. Under conditions of sulfur or magnesium deficiency, leaf chlorophyll content was less affected in transgenic plants, demonstrating the physiological relevance of shoot element accumulation. Our approach might contribute to improve drought tolerance, nutrient efficiency, and nutrient content of crop plants.

Summary Cytokinins are hormones that regulate many developmental and physiological processes in plants. Recent work has revealed that the cytokinin signal is transduced by two‐component systems to the nucleus where target genes are activated. Most of the rapid transcriptional responses are unknown. We measured immediate‐early and delayed cytokinin responses through genome‐wide expression profiling with the Affymetrix ATH1 full genome array (Affymetrix Inc., Santa Clara, CA, USA). Fifteen minutes after cytokinin treatment of 5‐day‐old Arabidopsis seedlings, 71 genes were upregulated and 11 genes were downregulated. Immediate‐early cytokinin response genes include a high portion of transcriptional regulators, among them six transcription factors that had previously not been linked to cytokinin. Five plastid transcripts were rapidly regulated as well, indicating a rapid transfer of the signal to plastids or direct perception of the cytokinin signal by plastids. After 2 h of cytokinin treatment genes coding for transcriptional regulators, signaling proteins, developmental and hormonal regulators, primary and secondary metabolism, energy generation and stress reactions were over‐represented. A significant number of the responding genes are known to regulate light (PHYA, PSK1, CIP8, PAT1, APRR), auxin (Aux/IAA), ethylene (ETR2, EIN3, ERFs/EREBPs), gibberellin (GAI, RGA1, GA20 oxidase), nitrate (NTR2, NIA) and sugar (STP1, SUS1) dependent processes, indicating intense crosstalk with environmental cues, other hormones and metabolites. Analysis of cytokinin‐deficient 35S:AtCKX1 transgenic seedlings has revealed additional, long‐lasting cytokinin‐sensitive changes of transcript abundance. Comparative overlay‐analysis with the software tool mapman identified previously unknown cytokinin‐sensitive metabolic genes, for example in the metabolism of trehalose‐6‐phosphate. Taken together, we present a genome‐wide view of changes in cytokinin‐responsive transcript abundance of genes that might be functionally relevant for the many biological processes that are governed by cytokinins.

This study on common marmosets Callithrix jacchus is the first to examine noise-dependent mechanisms of vocal plasticity in a New World monkey. Since acoustic communication can be considerably impaired by environmental noise, some animals have evolved adaptations to counteract its masking effects. The studied marmosets increased the sound level of their spontaneous calls in response to increased levels of white noise broadcast to them. Possibly, such noise-dependent adjustment of vocal amplitude serves to maintain a specific signal-to-noise ratio that is favourable for signal production. Concurrently, the adjustment of vocal amplitude can maintain a given active space for communication. In contrast to some bird species, no noiseinduced increase in the number of syllables per call series could be found, showing that an increased serial redundancy of vocal signals was not used to communicate under noisy conditions. Finally, we examined a possible noise-dependent prolongation of vocal signals. This approach was guided by the findings of perceptional studies, which suggest an increased detection probability of prolonged signals in noise by temporal summation. Marmosets indeed increased the duration of their call syllables along with increasing background noise levels. This is the first evidence of such mechanism of vocal plasticity in an animal communication system.

A survey of recent results is presented concerning the role of cytokinin degradation in plants, which is catalyzed by cytokinin oxidase/dehydrogenase (CKX) enzymes. An overview of Arabidopsis CKX gene expression suggests that their differential regulation by biotic and abiotic factors contributes significantly to functional specification. Here, we show using reporter gene and semiquantitative RT-PCR analyses regulation of individual CKX genes by cytokinin, auxin, ABA, and phosphate starvation. Partially overlapping expression domains of CKX genes and cytokinin-synthesizing IPT genes in meristematic tissues and endo-reduplicating cells lend support for a locally restricted function of cytokinin. On the other hand, their expression in vascular tissue suggests a function in controlling transported cytokinin. Recent studies led to a model for the biochemical reaction mechanism of CKX-mediated catalysis, which was refined on the basis of the three-dimensional enzyme structure. Last but not least, the developmental functions of CKX enzymes are addressed. The recent identification of the rice OSCKX2 gene as an important novel breeding tool is highlighted. Together the results corroborate the relevance of metabolic control in determining cytokinin activity.

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SUMMARYDegradation of the plant hormone cytokinin is catalyzed by cytokinin oxidase/dehydrogenase (CKX) enzymes. The Arabidopsis thaliana genome encodes seven CKX proteins which differ in subcellular localization and substrate specificity. Here we analyze the CKX7 gene, which to the best of our knowledge has not yet been studied. pCKX7:GUS expression was detected in the vasculature, the transmitting tissue and the mature embryo sac. A CKX7-GFP fusion protein localized to the cytosol, which is unique among all CKX family members. 35S:CKX7-expressing plants developed short, early terminating primary roots with smaller apical meristems, contrasting with plants overexpressing other CKX genes. The vascular bundles of 35S:CKX7 primary roots contained only protoxylem elements, thus resembling the wol mutant of the CRE1/AHK4 receptor gene. We show that CRE1/AHK4 activity is required to establish the CKX7 overexpression phenotype. Several cytokinin metabolites, in particular cis-zeatin (cZ) and N-glucoside cytokinins, were depleted stronger in 35S:CKX7 plants compared with plants overexpressing other CKX genes. Interestingly, enhanced protoxylem formation together with reduced primary root growth was also found in the cZ-deficient tRNA isopentenyltransferase mutant ipt2,9. However, different cytokinins were similarly efficient in suppressing 35S:CKX7 and ipt2,9 vascular phenotypes. Therefore, we hypothesize that the pool of cytosolic cytokinins is particularly relevant in the root procambium where it mediates the differentiation of vascular tissues through CRE1/AHK4. Taken together, the distinct consequences of CKX7 overexpression indicate that the cellular compartmentalization of cytokinin degradation and substrate preference of CKX isoforms are relevant parameters that define the activities of the hormone.

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