Signaling network regulating plant branching: Recent advances and new challenges.

Auxin alone or supplemented with cytokinins and strigolactones were long considered as the main player(s) in the control of apical dominance (AD) and correlative inhibition of the lateral bud outgrowth, the processes that shape the plant phenotype. However, past decade data indicate a more sophisticated pathways of AD regulation, with the involvement of mobile carbohydrates which perform both signal and trophic functions. Here we provide a critical comprehensive overview of the current status of the AD problem. This includes insight into intimate mechanisms regulating directed auxin transport in axillary buds with participation of phytohormones and sugars. Also roles of auxin, cytokinin and sugars in the dormancy or sustained growth of the lateral meristems were assigned. This review not only provides the latest data on implicated phytohormone crosstalk and its relationship with the signaling of sugars and abscisic acid, new AD players, but also focuses on the emerging biochemical mechanisms, at first positive feedback loops involving both sugars and hormones, that ensure the sustained bud growth. Data show that sugars act in concert with cytokinins but antagonistically to strigolactone signaling. A complex bud growth regulating network is demonstrated and unresolved issues regarding the hormone-carbohydrate regulation of AD are highlighted.

Auxin-cytokinin interactions in the regulation of correlative inhibition in two-branched pea seedlings.

A model system of 10-12 day-old, two-branched (2-B) pea (Pisum sativum L. cv. Adagumsky) seedlings was used to study the roles of endogenous auxin indole-3-acetic acid (IAA) and cytokinins (CKs) in the interaction between the shoots. The IAA export activity (IEA) from shoots was 2-fold higher in one-branched (1-B) plants with one shoot removed than in the 2-B plants, while tZ-type cytokinin contents in xylem sap were 4-fold greater in the 1-B plants than in 2-B plants. Exogenous 6-benzylaminopurine introduced into the vascular stream of one shoot enhanced its IEA. Therefore, xylem cytokinin appears to control both growth and IEA in branches. In the hypocotyls of 1-B and 2-B plants, IAA contents were equal in both cases, while the levels of tZ-type cytokinins were different. These data do not agree with the well-supported role of auxin in down-regulating CK content. The observed paradox may be explained by assuming that a steady-state IAA level in the hypocotyls is feedback regulated via xylem cytokinin, which controls the delivery of IAA from the shoots. As a result, the level of IAA in the hypocotyl is most likely maintained at a threshold below which a decrease in auxin content can switch on CK synthesis that will increase xylem cytokinin levels, thereby stabilizing the level of IAA in the hypocotyl. Therefore, our results suggest that correlative inhibition in the 2-B pea system is a function of an IAA/CK feedback loop, in which cytokinin essentially acts as a second messenger for IAA.