Plant development is governed by signaling molecules called phytohormones. Typically, in certain developmental processes more than 1 hormone is implicated and, thus, coordination of their overlapping activities is crucial for correct plant development. However, molecular mechanisms underlying the hormonal crosstalk are only poorly understood. Multiple hormones including cytokinin and auxin have been implicated in the regulation of root development. Here we dissect the roles of cytokinin in modulating growth of the primary root. We show that cytokinin effect on root elongation occurs through ethylene signaling whereas cytokinin effect on the root meristem size involves ethylene-independent modulation of transport-dependent asymmetric auxin distribution. Exogenous or endogenous modification of cytokinin levels and cytokinin signaling lead to specific changes in transcription of several auxin efflux carrier genes from the PIN family having a direct impact on auxin efflux from cultured cells and on auxin distribution in the root apex. We propose a novel model for cytokinin action in regulating root growth: Cytokinin influences cell-to-cell auxin transport by modification of expression of several auxin transport components and thus modulates auxin distribution important for regulation of activity and size of the root meristem.

The versatile functionality and physiological importance of the phytohormone auxin is a major focus of attention in contemporary plant science. Recent studies have substantially contributed to our understanding of the molecular mechanisms underlying the physiological role of auxin in plant development. The mechanism of auxin action includes both fast responses not involving gene expression, possibly mediated by Auxin Binding Protein 1 (ABP1), and slower responses requiring auxin-regulated gene expression mediated by F-box proteins. These two mechanisms of action have been described to varying degrees for the major endogenous auxin indole-3-acetic acid (IAA) and for the synthetic auxins 2,4-dichlorophenoxyacetic acid (2,4-D) and naphthalene-1-acetic acid (NAA). However, in addition to IAA, plants synthesize three other compounds that are commonly regarded as "endogenous auxins", namely, 4-chloroindole-3-acetic acid (4-Cl-IAA), indole-3-butyric acid (IBA) and phenylacetic acid (PAA). Although a spectrum of auxinic effects has been identified for all these as well as several other endogenous compounds, we remain largely ignorant of many aspects of their mechanisms of action and the extent to which they contribute to auxin-regulated plant development. Here, we briefly summarize the action of IBA, 4-Cl-IAA and PAA, and discuss the extent to which their action overlaps with that of IAA or results from their metabolic conversions to IAA. Other possible pathways for their action are considered. We present a scheme for homeostatic regulation of IAA levels that embraces other endogenous auxins in terms of the described mechanism of auxin action including its receptor and downstream signal transduction events.

Auxin is a key coordinative signal required for many aspects of plant development and its levels are controlled by auxin metabolism and intercellular auxin transport. Here we find that a member of PIN auxin transporter family, PIN8 is expressed in male gametophyte of Arabidopsis thaliana and has a crucial role in pollen development and functionality. Ectopic expression in sporophytic tissues establishes a role of PIN8 in regulating auxin homoeostasis and metabolism. PIN8 co-localizes with PIN5 to the endoplasmic reticulum (ER) where it acts as an auxin transporter. Genetic analyses reveal an antagonistic action of PIN5 and PIN8 in the regulation of intracellular auxin homoeostasis and gametophyte as well as sporophyte development. Our results reveal a role of the auxin transport in male gametophyte development in which the distinct actions of ER-localized PIN transporters regulate cellular auxin homoeostasis and maintain the auxin levels optimal for pollen development and pollen tube growth.