Artificial selection reveals complex genetic architecture of shoot branching and its response to nitrate supply in Arabidopsis.

Quantitative traits may be controlled by many loci, many alleles at each locus, and subject to genotype-by-environment interactions, making them difficult to map. One example of such a complex trait is shoot branching in the model plant Arabidopsis, and its plasticity in response to nitrate. Here, we use artificial selection under contrasting nitrate supplies to dissect the genetic architecture of this complex trait, where loci identified by association mapping failed to explain heritability estimates. We found a consistent response to selection for high branching, with correlated responses in other traits such as plasticity and flowering time. Genome-wide scans for selection and simulations suggest that at least tens of loci control this trait, with a distinct genetic architecture between low and high nitrate treatments. While signals of selection could be detected in the populations selected for high branching on low nitrate, there was very little overlap in the regions selected in three independent populations. Thus the regulatory network controlling shoot branching can be tuned in different ways to give similar phenotypes.

Strigolactones and the control of plant development: lessons from shoot branching.

Strigolactones (SLs) were originally identified through their activities as root exudates in the rhizosphere; however, it is now clear that they have many endogenous signalling roles in plants. In this review we discuss recent progress in understanding SL action in planta, particularly in the context of the regulation of shoot branching, one of the best-characterized endogenous roles for SLs. Rapid progress has been made in understanding SL biosynthesis, but many questions remain unanswered. There are hints of as yet unidentified sources of SL, as well as unknown SL-like molecules with important signalling functions. SL signalling is even more enigmatic. Although a likely receptor has been identified, along with some candidate immediate downstream targets, our understanding of how these targets mediate SL signalling is limited. There is still considerable uncertainty about whether the targets of SL signalling are primarily transcriptional or not. There is at least one non-transcriptional target, because a rapid primary response to SL is the removal of PIN1 auxin exporter proteins from the plasma membrane in vascular-associated cells of the stem. We discuss how the various early events in SL signalling could result in the observed changes in shoot branching.

Auxin and strigolactone signaling are required for modulation of Arabidopsis shoot branching by nitrogen supply.

The degree of shoot branching is strongly affected by environmental conditions, such as nutrient availability. Here we demonstrate that nitrate limitation reduces shoot branching in Arabidopsis (Arabidopsis thaliana) both by delaying axillary bud activation and by attenuating the basipetal sequence of bud activation that is triggered following floral transition. Ammonium supply has similar effects, suggesting that they are caused by plant nitrogen (N) status, rather than direct nitrate signaling. We identify increased auxin export from active shoot apices, resulting in increased auxin in the polar auxin transport stream of the main stem, as a likely cause for the suppression of basal branches. Consistent with this idea, in the auxin response mutant axr1 and the strigolactone biosynthesis mutant more axillary growth1, increased retention of basal branches on low N is associated with a failure to increase auxin in the main stem. The complex interactions between the hormones that regulate branching make it difficult to rule out other mechanisms of N action, such as up-regulation of strigolactone synthesis. However, the proposed increase in auxin export from active buds can also explain how reduced shoot branching is achieved without compromising root growth, leading to the characteristic shift in relative biomass allocation to the root when N is limiting.