The maize mutant barren stalk1 is defective in axillary meristem development
Abstract
barren stalk1 is a recessive mutant of maize that has no tassel branches, spikelets, tillers, or ears. Here we present a detailed characterization of the ba1 mutant phenotype, including scanning electron microscopy of developing inflorescences, in situ hybridization analysis using a meristem marker, molecular mapping, and genetic analysis demonstrating an epistatic relationship between ba1 and teosinte branched1 (tb1). These data show that the primary defect in the ba1 mutant is a failure in axillary meristem development.
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ROX in Arabidopsis, LAX1 in rice, and BA1 in maize influence axillary meristem development.
“Another factor is a basic helix-loop-helix (bHLH) protein called REGULATOR OF AXILLARY MERISTEM FORMATION (ROX) in Arabidopsis (Yang et al., 2012), LAX PANICLE1 (LAX1) in rice (Komatsu et al., 2001, 2003) and BARREN STALK1 (BA1) in maize (Ritter et al., 2002; Gallavotti et al., 2004).”
From Shaping plant architecture (2015) · cites this paper as Ritter et al., 2002
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BackgroundThe role of barren stalk1 in the architecture of maize
Gallavotti, A., Zhao, Q., Kyozuka, J., Meeley, R. B., Ritter, M. K., Doebley, J. F., Enrico Pè, M., & Schmidt, R. J. (2004) · Nature · doi:10.1038/nature03148 · Cited by 365 · see citing works
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“In maize (Zea mays), barren stalk1 (ba1) and barren inflorescence2 (bif2) mutants exhibit severe suppression of all types of AM (McSteen and Hake, 2001; Ritter et al., 2002; Gallavotti et al., 2004).”
BackgroundTwo-Step Regulation of LAX PANICLE1 Protein Accumulation in Axillary Meristem Formation in Rice
Oikawa, T., & Kyozuka, J. (2009) · The Plant Cell · doi:10.1105/tpc.108.065425 · Cited by 251 · see citing works
“In the mutants of the ba1 gene, an ortholog of LAX1, AM formation is severely suppressed, while subtending bracts are produced normally (Ritter et al., 2002; Gallavotti et al., 2004).”
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“…factor (Bortiri et al., 2006) that controls the expression of RA1, a C2H2 RESEARCH ARTICLE Development 137 (8) D E V E LO P M E N T zinc-finger transcription factor (Vollbrecht et al., 2005). ra3 encodes a trehalose-6-phosphate phosphatase that may also act upstream of ra1 (Satoh et al., 2006).”
Teichmann, T., & Muhr, M. (2015) · Frontiers in Plant Science · doi:10.3389/fpls.2015.00233 · Cited by 228 · see citing works
“Another factor is a basic helix-loop-helix (bHLH) protein called REGULATOROF AXILLARYMERISTEM FORMATION (ROX) inArabidopsis (Yang et al., 2012), LAXPANICLE1 (LAX1) in rice (Komatsu et al., 2001, 2003) and BARREN STALK1 (BA1) in maize (Ritter et al., 2002; Gallavotti et al., 2004).”
BackgroundMolecular Control of Grass Inflorescence Development
Zhang, D., & Yuan, Z. (2014) · Annual Review of Plant Biology · doi:10.1146/annurev-arplant-050213-040104 · Cited by 235 · see citing works
“These phenotypes in ba1 are similar to those of the bif2 mutant (97, 119, 127).”
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“…et al., 2007), and BA1 encodes a basic helix-loop-helix transcription factor (Gallavotti et al., 2004). ba1 mutants completely lack all axillary meristems, resulting in a complete sterile plant essentially composed of a stalk and several leaves (Ritter et al., 2002; Gallavotti et al., 2004).”
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McSteen, P. (2009) · Plant Physiology · doi:10.1104/pp.108.129056 · Cited by 193 · see citing works
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BackgroundAuxin and Monocot DevelopmentPreprint
McSteen, P. (2010) · Cold Spring Harbor Perspectives in Biology · doi:10.1101/cshperspect.a001479 · Cited by 182 · see citing works
“…has not yet been reported (Komatsu et al. 2003; Gallavotti et al. 2004). ba1 mutants in maize and lax1 mutants in rice do not produce axillary meristems during inflorescence development but the bract leaves that subtend axillary meristems are unaffected (Ritter et al. 2002; Komatsu et al. 2003).”
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Gallavotti, A. (2013) · Journal of Experimental Botany · doi:10.1093/jxb/ert141 · Cited by 177 · see citing works
“this model is the observation that in ba1 tassels the suppressed bracts seem indeed enlarged relative to those present in normal tassels (Ritter et al., 2002; Gallavotti et al., 2008b).”
ResultMcSteen, P., Malcomber, S., Skirpan, A., Lunde, C., Wu, X., Kellogg, E., & Hake, S. (2007) · Plant Physiology · doi:10.1104/pp.107.098558 · Cited by 186 · see citing works
“…panicle1 (lax1) are required for initiation of axillary meristems during both vegetative and inflorescence development (Komatsu et al., 2001, 2003a; Ritter et al., 2002; Gallavotti et al., 2004). ba1/lax1 are members of a grass-specific basic helix-loop-helix gene family, suggesting that some of…”
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“…in maize inflorescence, such as barren stalk1 (Ritter et al., 2002; Gallavotti et al., 2004), ramosa1 (Vollbrecht et al., 2005), ramosa2 (Bortiri et al., 2006), ramosa3 (Satoh-Nagasawa et al., 2006), and suppressor of sessile spikelets1 (Doebley et al., 1995), should be taken into account.”
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“Another gene that similarly affects both vegetative and inflorescence axillary meristem initiation in maize is barren stalk1 (Zm-ba1) (Ritter et al., 2002; Gallavotti et al., 2004).”
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“Similarly, maize ba1mutants are compromised in lateral meristem initiation at all stages of development (Ritter et al., 2002; Gallavotti et al., 2004).”
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“was isolated in 1928 (Hofmeyer, 1930; Ritter et al., 2002 ).”
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“Another gene that similarly affects both vegetative and inflorescence axillary meristem initiation is Barren stalk1 (Ba1) in maize [32,33], although its rice equivalent, Lax panicle1 (Lax1), only affects inflorescence branching [34].”
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“Similarly, maize mutants with the ba1 allele fail to initiate lateral meristems during both vegetative and reproductive development, resulting in unbranched, short, and sterile tassels without spikelets (Ritter et al., 2002; Gallavotti et al., 2004; Skirpan et al., 2008; Yao et al., 2019).”
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“The ba1 mutants also fail to initiate BMs and SPMs in maize (Ritter et al., 2002; Gallavotti et al., 2004).”
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“Genetic experiments using single and double mutants have also shown that ba1 is completely epistatic to tb1 (Ritter et al. 2002; Gallavotti et al. 2004).”
MethodDevelopment of the Inflorescences
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Self-citationThe barren stalk2 Gene Is Required for Axillary Meristem Development in Maize
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“Similarly, the maize gene BARREN STALK 1 (BA1), affects tiller and tassel development [21].”
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“... basis of whether they affect meristem initiation (maize barren stalk1 [ba1], tomato lateral suppressor [ls], A. thaliana lateral suppressor [las], tomato blind [bl], and rice lax panicle [lax] and monoculm1 [moc1]), meristem outgrowth (A. thaliana more axillary growth [max], pea ramosus [rms], teosinte teosinte br…”
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Boundary domain genes were recruited to suppress bract growth and promote branching in maize
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“…2001;McSteen et al., 2007; Barazesh and McSteen, 2008a). barren stalk1 (ba1), which is proposed to act both upstream and downstream of auxin transport, also produces single spikelets when mutated (Ritter et al., 2002; Gallavotti et al., 2004, 2008; Wu and McSteen, 2007; Skirpan et al., 2008).”
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“These regular bulges are bract primordia whose outgrowth is usually inhibited during normal tassel development (Ritter et al. 2002; Gallavotti et al. 2004).”
The Ha‐ROXL gene is required for initiation of axillary and floral meristems in sunflower
Basile, A., Fambrini, M., Tani, C., Shukla, V., Licausi, F., & Pugliesi, C. (2019) · genesis · doi:10.1002/dvg.23307 · Cited by 16 · see citing works
“As SAM activity products, it is possible to distinguish metameric units, named phytomers that are composed of a node, an internode, a leaf and an AM (Rutishauser & Sattler, 1985).”
BackgroundKRN5b regulates maize kernel row number through mediating phosphoinositol signalling
Shen, X., Liu, L., Tran, T., Ning, Q., Li, M., Huang, L., Zhao, R., Li, Y., Qing, X., Jackson, D. P., et al. (2024) · Plant Biotechnology Journal · doi:10.1111/pbi.14463 · Cited by 16 · see citing works
“…(i.e., few branches and spikelets) of the mutants with abnormal auxin biosynthesis, transport, and signalling, including barren inflorescence2 ( bif2 ) and barren stalk2 ( ba2 ) (Gallavotti et al ., 2011; McSteen and Hake, 2001; Ritter et al ., 2002; Tabuchi et al ., 2011; Yao et al ., 2019).”
Chen, M.-S., Zhao, M., Wang, G., et al. (2019) · BMC Plant Biology · doi:10.1186/s12870-019-2069-3 · Cited by 15 · see citing works
Orchid biotechnology for Indonesian orchids conservation and industry
Semiarti, E. (2018) · AIP conference proceedings · doi:10.1063/1.5050118 · Cited by 15 · see citing works
Ahmad, S., Chen, J., Chen, G.-Z., et al. (2022) · Frontiers in Plant Science · doi:10.3389/fpls.2022.942591 · Cited by 14 · see citing works
Barazesh, S., Nowbakht, C., & McSteen, P. (2009) · Genetics · doi:10.1534/genetics.108.099390 · Cited by 13 · see citing works
“…barren stalk1 (ba1) produce fewer branches and spikelets in the inflorescence due to defects in axillary meristem initiation (McSteen and Hake 2001; Ritter et al. 2002; Barazesh and McSteen 2008a; Gallavotti et al. 2008). spi1 functions in localized auxin biosynthesis, while bif1 and bif2 regulate…”
BackgroundInfluentialThe Secrets of Meristems Initiation: Axillary Meristem Initiation and Floral Meristem Initiation
Yang, Q., Yuan, C., Cong, T., & Zhang, Q. (2023) · Plants · doi:10.3390/plants12091879 · Cited by 11 · see citing works
“The AM and inflorescence branching cannot develop normally in the ba1 mutant, and, similar to the LAX1 and LAX2 protein, the BA2 protein could also form heterodimerization with BA1 [46–48].”
BackgroundLv, Y., Zhang, X., Hu, Y., Liu, S., Yin, Y., & Wang, X. (2023) · Frontiers in Plant Science · doi:10.3389/fpls.2023.1162828 · Cited by 10 · see citing works
“The phenotypes of ba1 are similar to the bif2 mutant (Mcsteen and Hake, 2001; Ritter et al., 2002; Skirpan et al., 2008).”
BackgroundIdentification of QTL for fasciated ear related traits in maize
Mei, X., Dong, E., Liang, Q., et al. (2020) · Crop Science · doi:10.1002/csc2.20435 · Cited by 10 · see citing works
Phillips, K. A., Skirpan, A. L., Kaplinsky, N. J., & McSteen, P. (2009) · American Journal of Botany · doi:10.3732/ajb.0800268 · Cited by 10 · see citing works
Ni, J., You, C., Chen, Z., Tang, D., Wu, H., Deng, W., Wang, X., Yang, J., Bao, R., Liu, Z., et al. (2023) · Molecular Breeding · doi:10.1007/s11032-023-01431-y · Cited by 9 · see citing works
“Barren stalk1 ( ba1 ) encodes a basic helix-loop-helix transcription factor that regulates the initiation of AMs, and the ba1 mutants are usually unbranched and shortened (Gal-lavotti et al. 2004; Ritter et al. 2002) .”
BackgroundGenetic and Hormonal Regulation of Maize Inflorescence Development
Thompson, B. E. (2014) · Advances in botanical research · doi:10.1016/b978-0-12-417162-6.00009-2 · Cited by 9 · see citing works
Kyozuka, J. (2014) · Advances in botanical research · doi:10.1016/b978-0-12-417162-6.00007-9 · Cited by 8 · see citing works
Nucleotide Diversity of the Maize ZmCNR13 Gene and Association With Ear Traits
Zuo, Z., Lu, Y., Zhu, M., et al. (2021) · Frontiers in Genetics · doi:10.3389/fgene.2021.773597 · Cited by 5 · see citing works
“, 2005), bif2 (McSteen and Hake 2001) and ba1 (Ritter et al., 2002) for tassel morphology.”
Li, W. (2012) · doi:10.31274/etd-180810-1038 · Cited by 5 · see citing works
“The expression of tb1 is conserved in husk leaves of maize and teosinte [10].”
BackgroundInfluentialReview: Recent advances in unraveling the genetic architecture of kernel row number in maize
Wang, Y., Shaw, R. K., & Fan, X. (2024) · Plant Science · doi:10.1016/j.plantsci.2024.112366 · Cited by 4 · see citing works
Li, J., Meng, D., Yu, H., et al. (2019) · Genetic Resources and Crop Evolution · doi:10.1007/s10722-019-00805-6 · Cited by 4 · see citing works
Wang, H., Liu, S., Ma, S., Wang, Y., Yang, H., Liu, J., Li, M., Cui, X., Liang, S., Cheng, Q., et al. (2023) · International Journal of Molecular Sciences · doi:10.3390/ijms241612718 · Cited by 3 · see citing works
“In maize, the BA1 gene, a rice LAX homolog, regulates AM formation and maintenance, and tillering is lost in the mutant [93].”
BackgroundGenetics of Whole Plant Morphology and Architecture
Rossini, L., Muehlbauer, G. J., Okagaki, R. J., Salvi, S., & von Korff, M. (2018) · Compendium of plant genomes · doi:10.1007/978-3-319-92528-8_13 · Cited by 3 · see citing works
Study of RAMOSA1 function during maize inflorescence development
Yang, X. (2011) · doi:10.31274/etd-180810-1469 · Cited by 3 · see citing works
Transcriptomic and metabolic changes during tassel branching development in maize
Tai, Y., Lyu, X., Pan, F., Meng, L., Cheng, Z., Xu, Z., Li, M., Hao, Z., Zhang, D., Yong, H., et al. (2025) · BMC Plant Biology · doi:10.1186/s12870-025-06607-6 · Cited by 2 · see citing works
“BA1 encodes a noncanonical basic helix-loop-helix protein that controls the auxin-mediated initiation of axillary meristems, and the barren stalk 1 ( ba1 ) mutant lacks tassel branches, indicating BA1 promotes the formation of branch mer-istems [16].”
Dynamic expression of POH1 gene in shoot development during in vitro culture of Phalaenopsis orchid
Semiarti, E., Slamet, A., Rizal, R., et al. (2016) · AIP conference proceedings · doi:10.1063/1.4953493 · Cited by 2 · see citing works
Cereal inflorescence: features of morphology, development and genetic regulation of morphogenesis
Dobrovolskaya, O. B., & Dresvyannikova, A. E. (2018) · Vavilov Journal of Genetics and Breeding · doi:10.18699/vj18.420 · Cited by 2 · see citing works
Genetic analysis of ear development and tassel architecture in maize (Zea mays L. ssp. mays)
Mayor, M. L. (2008) · doi:10.31274/etd-180810-1831 · Cited by 2 · see citing works
“The loci Suppressor of sessile spikelet (Sos1), barren stalk1 (ba1) and barren inflorescence2 (bif2) together with tb1 affect the initiation and maintenance of inflorescence meristem (Gallavotti et al. 2004; McSteen and Hake 2001; Ritter et al. 2002).”
BackgroundMaize growth and yield as affected by different soil fertility regimes in a long term trial
Dlamini, J. C. (2015) · UpSpace Institutional Repository (University of Pretoria) · Cited by 2 · see citing works
Molecular cloning of the GhLOM and GhROX genes associated with axillary meristem formation in cotton
Dai, T. C., & Wang, Z. (2016) · Russian Journal of Plant Physiology · doi:10.1134/s1021443716030031 · Cited by 1 · see citing works
Ma, T., Li, L., Zhao, Y., et al. (2021) · Biology · doi:10.3390/biology10050437 · Cited by 1 · see citing works
“Genetic studies have demonstrated that the TtBH/ Wheat FRIZZY PANICLE (WFZP) gene determines the formation of the lateral meristem in wheat, and this gene is orthologous to the AP2/EREBP-like transcription factor genes responsible for branched silkless1 (BD1) in maize and for frizzy panicle (FZP) in rice [10,11].”
BackgroundCharacterization of two novel barren mutants in maize.
Buck, A. (2013) · eScholarship (California Digital Library)
Hormonal Regulation of Branching in Grasses 1,2(C)
McSteen, P. (2009)
Haaning, A. (2018) · University of Minnesota Digital Conservancy (University of Minnesota)
Bhatt, A. M. (2017) · Annual Plant Reviews online · doi:10.1002/9781119312994.apr0168
Axillary Meristem Development in Plants
Zhang, C., Liu, L., Guo, S., & Duan, B. (2022) · Current Chinese Science · doi:10.2174/2210298102666220927100443
The roles of transcription factors in the development of plant meristems
Dong, Q., & Zhang, C. (2022) · Elsevier eBooks · doi:10.1016/b978-0-323-90613-5.00008-x
Genetics and development of pseudanthia in the genus Helianthus: A review
Rogo, U., Fambrini, M., Pugliesi, C., & Simoni, S. (2025) · Biologia · doi:10.1007/s11756-025-02024-0 · Cited by 1 · see citing works
“In ba1 , the result is flower-less tassels with enlarged bracts (Ritter et al. 2002), resembling a reminiscence of what was observed histologically in the floret primordia of the sunflower mf mutant.”
BS-D275: A novel locus on the chromosome arm 2DS governing branched spike development in wheat
Qu, Y., Ren, Y., Hu, J., Liu, Y., Qiu, D., Li, Y., Wu, Q., He, M., Guo, Q. Q., Sun, Y., et al. (2025) · Theoretical and Applied Genetics · doi:10.1007/s00122-025-05091-8
“…S2DG0331600 / TraesCS2D02G118200 , located at approximately 73.09 Mb), is the wheat ortholog of maize ( Zea mays L.) branched silkless1 ( BD1 ) (Ritter et al. 2002; Gallavotti et al. 2004; Poursarebani et al. 2015) and rice frizzy panicle Within the BS-D275 interval on chromosome arm 2DS,…”
InfluentialSu, Y. H., Tang, L. P., & Zhang, X. S. (2026) · Springer eBooks · doi:10.1007/978-981-95-7033-1_2
- Authors
- Ritter, M. K., Padilla, C. M., & Schmidt, R. J.
- Affiliations
- University of California San Diego (US)
- Corresponding author
- Schmidt, Robert J.
- Venue
- American Journal of Botany
- Publisher
- Wiley
- Published
- 2002-02
- Volume
- 89
- Issue
- 2
- Pages
- 203-210
- ISSN
- 0002-9122 (ISSN-L)
- 1537-2197
- OpenAlex
- W2130989933
- Cited by
- 90
- References
- 31
- Citations by year
- 2026: 1 · 2025: 3 · 2024: 3 · 2023: 5 · 2022: 5 · 2021: 4 · 2020: 1 · 2019: 5 · 2018: 4 · 2017: 4 · 2016: 2 · 2015: 5 · 2014: 4 · 2013: 3 · 2012: 2
- FWCI
- 3.37
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- American Journal of Botany (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Science Foundation
- Topics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Molecular Biology Research
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Genetic and Mutation Studies
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Parasitism and Resistance
- Keywords
- maize, inflorescence development, TEOSINTE BRANCHED1, epistasis, tillering, molecular mapping, recessive mutants
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