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  • Review Article
  • Published:

Signal integration in the control of shoot branching

Key Points

  • Auxins, cytokinins and strigolactones are three classes of hormones that regulate bud activation and thereby regulate shoot branching. These hormones are transported throughout the plant, forming a systemic network that allows the integration of information between different plant organs and with the environment.

  • The identification and characterization in various plant species of a class of highly branching mutants that lack, or are impaired in the perception of, the acropetally transported hormones known as strigolactones was central to recent progress in understanding the mechanisms of shoot branching control.

  • One model for bud activation is based on the auxin transport canalization-based model, which stipulates that an initial flux from an auxin source to an auxin sink is gradually canalized into files of cells with high levels of highly polarized transporters. The process of canalization is driven by a positive feedback loop in which auxin flux upregulates and polarizes auxin efflux facilitators in the direction of the auxin flow, resulting in the formation of auxin transport canals.

  • In the auxin transport canalization-based model for bud activation, buds are considered as auxin sources, and the stem acts as an auxin sink owing to its ability to transport auxin away to the root. Buds must export auxin to be activated, and upon bud activation, auxin transport from active buds reduces the sink strength of the stem, and thus prevents other buds from exporting their auxin.

  • According to an alternative model for bud activation, known as the second messenger model, auxin derived from the shoot apex inhibits bud outgrowth by regulating the production of a second messenger, which moves directly into the bud to control its activity. Cytokinins and strigolactones are two candidate hormones to serve as second messengers.

  • Environmental inputs, such as altered light quality or nutrient deficiency in the soil, affect shoot branching using a network of interacting hormones including auxin, cytokinins and strigolactones. The detailed molecular mechanisms of the integration of the environmental signals into the system remain to be unravelled.

Abstract

Shoot branching is a highly plastic developmental process in which axillary buds are formed in the axil of each leaf and may subsequently be activated to give branches. Three classes of plant hormones, auxins, cytokinins and strigolactones (or strigolactone derivatives) are central to the control of bud activation. These hormones move throughout the plant forming a network of systemic signals. The past decade brought great progress in understanding the mechanisms of shoot branching control. Biological and computational studies have led to the proposal of two models, the auxin transport canalization-based model and the second messenger model, which provide mechanistic explanations for apical dominance.

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Figure 1: A schematic of plant shoot architecture.
Figure 2: Auxin and strigolactones inhibit bud outgrowth.
Figure 3: The long-distance hormonal network in shoot branching control.
Figure 4: Auxin transport canalization and bud activation.
Figure 5: The strigolactone pathway.
Figure 6: The second messenger model for bud activation.
Figure 7: The effect of the environment on bud activation.

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Acknowledgements

We thank S. Day for critical reading of the manuscript. Work in O.L.'s group is funded by the Biotechnology and Biological Science Council, The European Commission and the Gatsby Foundation.

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Glossary

Meristem

A group of cells with stem-cell-like properties. These cells continuously divide to build the plant body, while also maintaining a pool of pluripotent cells.

Axil

The region directly above the point where the leaf joins the stem.

Shoot apex

The shoot tip, the topmost part of the shoot axis that contains the apical meristem.

Basipetal transport

Movement away from the apex.

Polar auxin transport

Active and directional transport of auxin.

Acropetal transport

Movement towards the apex.

Xylem

Vascular tissue that delivers water, hormones and mineral nutrients from the root to the shoot.

Phyllotactic

Relating to the initiation of leaves.

Bistable

A dynamic system that has two stable states, with intermediate states being unstable and thus rapidly resolving towards one or other of the stable states.

Far-red light

Wavelengths of light of around 700–800 nm. Red light is absorbed for photosynthesis, but far-red light is not, so a decrease in the ratio of red light to far-red light is indicative of shading by another plant.

Phytochrome

A protein belonging to a class of plant photoreceptors that are mainly responsible for perception of red and far red light.

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Domagalska, M., Leyser, O. Signal integration in the control of shoot branching. Nat Rev Mol Cell Biol 12, 211–221 (2011). https://doi.org/10.1038/nrm3088

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