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Correction ARTICLE

Front. Plant Sci., 09 May 2019 |

Corrigendum: miR156/SPL10 Modulates Lateral Root Development, Branching and Leaf Morphology in Arabidopsis by Silencing AGAMOUS-LIKE 79

Ruimin Gao1, Ying Wang1, Margaret Y. Gruber2 and Abdelali Hannoufa1*
  • 1London Research and Development Center, Agriculture and Agri-Food Canada, London, ON, Canada
  • 2Saskatoon Research and Development Center, Agriculture and Agri-Food Canada, Saskatoon, SK, Canada

A Corrigendum on
miR156/SPL10 Modulates Lateral Root Development, Branching and Leaf Morphology in Arabidopsis by Silencing AGAMOUS-LIKE 79

by Gao, R., Wang, Y., Gruber, M. Y., and Hannoufa, A. (2018). Front. Plant Sci. 8:2226. doi: 10.3389/fpls.2017.02226

In the original article, there was a mistake in the legend for Figure 4D, Figure 8A as published. There was an error in our CRISPR-generated AGL79KD mutant. In the Results, subsection Characterization of the AGL79KD Arabidopsis Mutant, we showed Arabidopsis knockdown mutants with a CRISPR-mutated AGL79. In performing follow up research we discovered that the plants had no point mutations as claimed in the original manuscript. As the researcher who carried out this experiment left the lab about 18 months ago, we reviewed the original sequence traces, and found that when reading the sequence in one direction (as the researcher probably did) it seemed to indicate point mutations, but when we sequenced in the reverse direction we could find no such point mutations in the AGL79 locus, and determined that this was indeed an error in reading the sequencing data. Figure legends that included this mutant have been modified to omit ALG79KD (Figures 4D and 8A).


Figure 4. Phenotypic characterization of AGL79 misexpression Arabidopsis plants. All the used plants were grown at the same time and conditions, and the comparisons were carried out when WT reached the bolting stager. (A) WT plants. (B) Arabidopsis plants with highest AGL79 gene over expression (OE). (C) Phenotypic display of SPL10 overexpression line (6mSPL10) (bar = 1.1 cm).


Figure 8. Complementary experiment to investigate relationship between AGL79 and SPL10. WT phenotype was observed when crossing (AGL79 overexpression plant and spl2spl10 double mutant. Bar = 1.5cm.

Due to the same error detailed above, there was a mistake in Figures 6, 7, Supplementary Figure 1 and the Supplementary Material Document 2. Figures and their corresponding legends that were specific to AGL79KD have been removed (Figures 6, 7, Supplementary Figure 1, and Supplementary Material Document 2). Figures have also been renumbered to reflect these changes. In addition, primers related to the generation and validation of CRISPR plants have been removed from Supplementary Table 2.

Furthermore, in the original article, the following eight references, that are related to AGL79 KD mutant descriptions/discussions, have been removed.

1. An, H., Roussot, C., Suarez-Lopez, P., Corbesier, L., Vincent, C., Pineiro, M., et al. (2004). CONSTANS acts in the phloem to regulate a systemic signal that induces photoperiodic flowering of Arabidopsis. Development 131, 3615–3626. doi: 10.1242/dev.01231

2. Bennett, T., Sieberer, T., Willett, B., Booker, J., Luschnig, C., and Leyser, O. (2006). The Arabidopsis MAX pathway controls shoot branching by regulating auxin transport. Curr. Biol. 16, 553–563. doi: 10.1016/j.cub.2006.01.058

3. Lee, J. H., Yoo, S. J., Park, S. H., Hwang, I., Lee, J. S., and Ahn, J. H. (2007). Role of SVP in the control of flowering time by ambient temperature in Arabidopsis. Genes Dev. 21, 397–402. doi: 10.1101/gad.1518407

4. Lei, Y., Lu, L., Liu, H. Y., Li, S., Xing, F., and Chen, L. L. (2014). CRISPR-P: a web tool for synthetic single-guide RNA design of CRISPR-system in plants. Mol. Plant 7, 1494–1496. doi: 10.1093/mp/ssu044

5. Michaels, S. D., Himelblau, E., Kim, S. Y., Schomburg, F. M., and Amasino, R. M. (2005). Integration of flowering signals in winter-annual Arabidopsis. Plant Physiol. 137, 149–156. doi: 10.1104/pp.104.052811

6. Ongaro, V., Bainbridge, K., Williamson, L., and Leyser, O. (2008). Interactions between axillary branches of Arabidopsis. Mol. Plant 1, 388–400. doi: 10.1093/mp/ssn007

7. Rameau, C., Bertheloot, J., Leduc, N., Andrieu, B., Foucher, F., and Sakr, S. (2014). Multiple pathways regulate shoot branching. Front. Plant Sci. 5:741. doi: 10.3389/fpls.2014.00741

8. Takada, S., and Goto, K. (2003). Terminal flower2, an Arabidopsis homolog of heterochromatin protein1, counteracts the activation of flowering locus T by constans in the vascular tissues of leaves to regulate flowering time. Plant Cell 15, 2856–2865. doi: 10.1105/tpc.016345

Lastly, corrections have been made to the Methods, Results, and Discussion of the original article and the AGL79KD mutant has been removed as detailed below.

In the Methods, subsection CRISPR-Cas9 Design and Screening for AGL79 Gene Editing has been removed.

Subsection Characterization of AGL79 Overexpression Plants has been corrected to:

“To investigate the role of AGL79 in Arabidopsis development, we generated transgenic plants with enhanced expression of AGL79. Compared to WT, the highest AGL79 overexpression plants flowered early and had fewer and smaller rosette leaves [Figures 4B, 5B (WT and Group 1)] much like the SPL10 overexpression plants (6mSLP10) (Figure 4C, Supplementary Figure 1). Transgenic Arabidopsis plants harboring the AGL79 overexpression construct were divided into three groups depending on AGL79 expression. Group 1 (lines L1, L2, and L3) had the highest AGL79 transcript levels in both the leaf and root tissues (Figure 5A), with lower expression in roots relative to leaves. Group 2 (lines L11, L12, and L20) had intermediate AGL79 expression, with variable expression levels between leaf and root (Figure 5A). Group 3 (lines L16, L18, and L27) displayed the lowest AGL79 gene transcripts, and there were no obvious differences in AGL79 transcript levels between the leaf and root (Figure 5A). Different phenotypes could be observed in these AGL79 overexpression plants depending on AGL79 expression levels (Figure 5B). Compared to WT (3 weeks after seed germination), Group 1 plants displayed fewer rosette leaves and early flowering time (Figure 5B). Group 2 plants displayed a phenotype similar to WT (Figure 5B). Group 3 plants showed more lateral shoot branches and a higher number of rosette leaves, as well as a significant delay in flowering (Figure 5B). In addition, the transcript level of SPL10 gene was also investigated in both the leaves and roots of the above-mentioned plants. Although changes in SPL10 expression could be detected in three groups of AGL79OE plants (Figure 5C), these changes did not follow any consistent trend, as found for AGL79 (Figure 5C), suggesting that AGL79 could be a downstream gene regulated by SPL10, and hence fluctuations in AGL79 expression would not affect the expression of the upstream SPL10 gene.”

Subsection Characterization of the AGL79KD Arabidopsis Mutant, has been removed.

Subsection Regulatory Relationship between AGL79 and SPL10, has been corrected to:

“As all the evidence derived from molecular and biological analysis (Figures 2A,B, 4C) revealed that AGL79 is likely regulated through the miR156-SPL pathway, we investigated whether a linear regulatory relationship exists between SPL10 and AGL79. Crossing AGL79OE plants and spl2spl10 double mutant produced F1 progeny showing WT-like phenotype (Figure 8B). The selected genotyping results of the double mutant (spl2spl10) and AGL79 OE plants are shown in Supplementary Figure 3. These results suggest a direct linear relationship between AGL79 and SPL10 genes.”

In the Discussion, paragraph two has been corrected to:

“The discovery that AGL79 is regulated by SPL10 may provide insight into how the latter regulates lateral root development in Arabidopsis (Yu et al., 2015). Currently lateral root formation in Arabidopsis is known to be regulated by two related AUXIN RESPONSE FACTORS (ARF7 and ARF19) via direct activation of LATERAL ORGAN BOUNDARIES DOMAIN and ASYMMETRIC LEAVES-LIKE (LBD/ASLs) (Okushima et al., 2007). In addition, lateral root formation in Arabidopsis is also redundantly regulated by cytokinin biosynthesis genes IPT3 and IPT5 and all three cytokinin histidine kinase receptor genes (AHK2, AHK3, and CRE1/AHK4) (Chang et al., 2013). The plant hormones (auxin, cytokinins, gibberellins, abscisic acid, ethylene, jasmonic acid, strigolactones, brassinosteroids, and salicylic acid) also regulate normal root growth and mediate root morphological responses to abiotic stress (Chang et al., 2013). Morphological analysis of Arabidopsis plants with enhanced expression of AGL79 revealed AGL79 to be involved in controlling shoot branching.”

and paragraph four should be removed and the last paragraph has been corrected to:

“In summary, our results suggest that the miR156/SPL10 regulatory pathway is involved in regulating plant lateral root growth by directly targeting and activating the expression of AGL79. By investigating the gain- of function of AGL79 transgenic plants, we also found AGL79 to be involved in regulating plant leaf shape, shoot branching, and flowering time. Further characterization of the AGL79 gene in other plant species, especially in major crops, will determine how conserved AGL79 is in plants. It can also be tested in crop improvement efforts to enhance resilience and productivity.”

The authors apologize for these errors and state that this does not change the scientific conclusions of the article in any way. The original article has been updated.

Supplementary Material

The Supplementary Material for this article can be found online at:


Chang, L., Ramireddy, E., and Schmulling, T. (2013). Lateral root formation and growth of Arabidopsis is redundantly regulated by cytokinin metabolism and signalling genes. J. Exp. Bot. 64, 5021–5032. doi: 10.1093/jxb/ert291

PubMed Abstract | CrossRef Full Text | Google Scholar

Okushima, Y., Fukaki, H., Onoda, M., Theologis, A., and Tasaka, M. (2007). ARF7 and ARF19 regulate lateral root formation via direct activation of LBD/ASL genes in Arabidopsis. Plant Cell 19, 118–130. doi: 10.1105/tpc.106.047761

PubMed Abstract | CrossRef Full Text | Google Scholar

Yu, N., Niu, Q. W., Ng, K. H., and Chua, N. H. (2015). The role of miR156/SPLs modules in Arabidopsis lateral root development. Plant J. 83, 673–685. doi: 10.1111/tpj.12919

PubMed Abstract | CrossRef Full Text | Google Scholar

Keywords: Arabidopsis, miR156, SPL10, lateral root, AGL79, flowering time, leaf morphology

Citation: Gao R, Wang Y, Gruber MY and Hannoufa A (2019) Corrigendum: miR156/SPL10 Modulates Lateral Root Development, Branching and Leaf Morphology in Arabidopsis by Silencing AGAMOUS-LIKE 79. Front. Plant Sci. 10:515. doi: 10.3389/fpls.2019.00515

Received: 26 February 2019; Accepted: 03 April 2019;
Published: 09 May 2019.

Edited and reviewed by: Jin-Gui Chen, Oak Ridge National Laboratory (DOE), United States

Copyright © 2019 Gao, Wang, Gruber and Hannoufa. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Abdelali Hannoufa,

Present Address: Ying Wang, Department of Biology, Carleton University, Ottawa, ON, Canada