Root architecture was determined together with shoot parameters under well watered and drought conditions in the field in three soybean cultivars (A5409RG, Jackson and Prima 2000). Morphology parameters were used to classify the cultivars into different root phenotypes that could be important in conferring drought tolerance traits. A5409RG is a drought-sensitive cultivar with a shallow root phenotype and a root angle of <40°. In contrast, Jackson is a drought-escaping cultivar. It has a deep rooting phenotype with a root angle of >60°. Prima 2000 is an intermediate drought-tolerant cultivar with a root angle of 40°-60°. It has an intermediate root phenotype. Prima 2000 was the best performing cultivar under drought stress, having the greatest shoot biomass and grain yield under limited water availability. It had abundant root nodules even under drought conditions. A positive correlation was observed between nodule size, above-ground biomass and seed yield under well-watered and drought conditions. These findings demonstrate that root OPEN ACCESSAgronomy 2014, 4 419 system phenotyping using markers that are easy-to-apply under field conditions can be used to determine genotypic differences in drought tolerance in soybean. The strong association between root and nodule parameters and whole plant productivity demonstrates the potential application of simple root phenotypic markers in screening for drought tolerance in soybean.

Drought is considered to be a major threat to soybean production worldwide and yet our current understanding of the effects of drought on soybean productively is largely based on studies on above-ground traits. Although the roots and root nodules are important sensors of drought, the responses of these crucial organs and their drought tolerance features remain poorly characterized. The symbiotic interaction between soybean and rhizobia facilitates atmospheric nitrogen fixation, a process that provides essential nitrogen to support plant growth and development. Symbiotic nitrogen fixation is important for sustainable agriculture, as it sustains plant growth on nitrogen-poor soils and limits fertilizer use for crop nitrogen nutrition. Recent developments have been made in our understanding of the drought impact on soybean root architecture and nodule traits, as well as underpinning transcriptome, proteome and also emerging metabolome information, with a view to improve the selection of more drought-tolerant soybean cultivars and rhizobia in the future. We conclude that the direct screening of root and nodule traits in the field as well as identification of genes, proteins and also metabolites involved in such traits will be essential in order to gain a better understanding of the regulation of root architecture, bacteroid development and lifespan in relation to drought tolerance in soybean.

Drought is the single most important factor limiting soybean (Glycine max L. Merr.) yields in the field. The following study was therefore undertaken to identify phenotypic markers for enhanced drought tolerance in nodulated soybeans. Leaf and nodule parameters were compared in three genotypes: Prima 2000, glyphosate-resistant A5409RG and Jackson, which had similar shoot biomass and photosynthesis rates at the third trifoliate leaf stage under water-replete conditions. When water was withheld at the third trifoliate leaf stage, photosynthesis, nodule numbers, nodule biomass and symbiotic nitrogen fixation (SNF) were greatly decreased. Significant cultivar-drought interactions were observed wi th respect to photosynthesis, which also showed a strong posi tive correlation with nodule SNF, particularly under drought conditions. Prima leaves had high water use efficiencies and they also maintained high photosynthetic electron transport efficiencies under long term drought. Moreover, Prima had the highest shoot biomass under both water-replete and drought conditions. A-5409RG was the most drought-sensitive genotype showing early closure of stomata and rapid inhibition of photosynthesis in response to drought. In addition to classifying the genotypes in relation to drought tolerance, the results demonstrate that the ability to sustain shoot biomass under nitrogen limitation is an important parameter, which can be easily applied in germplasm screening for drought tolerance in soybean.

CLE peptides are potentially involved in nodule organ development and in the autoregulation of nodulation (AON), a systemic process that restricts nodule number. A genome-wide survey of CLE peptide genes in the soybean glycine max genome resulted in the identification of 39 GmCLE genes, the majority of which have not yet been annotated. qRT-PCR analysis indicated two different nodulation-related CLE expression patterns, one linked with nodule primordium development and a new one linked with nodule maturation. Moreover, two GmCLE gene pairs, encoding group-III CLE peptides that were previously shown to be involved in AON, had a transient expression pattern during nodule development, were induced by the essential nodulation hormone cytokinin, and one pair was also slightly induced by the addition of nitrate. Hence, our data support the hypothesis that group-III CLE peptides produced in the nodules are involved in primordium homeostasis and intertwined in activating AON, but not in sustaining it.