Abstract
RUBylation plays essential roles in plant growth and development through regulating Cullin-RING ubiquitin E3 ligase (CRL) activities and the CRL-mediated protein degradations. However, the function of RUBylation in regulating kernel development remains unclear. Through genetic and molecular analyses of a small kernel 501 (smk501) mutant in maize (Zea mays), we cloned the smk501 gene, revealed its molecular function, and defined its roles in RUBylation pathway and seed development. Smk501 encodes a RUBylation activating enzyme E1 subunit ZmECR1 (E1 C-TERMINAL RELATED 1) protein. Destruction in RUBylation by smk501 mutation resulted in less embryo and endosperm cell number and smaller kernel size. The transcriptome and proteome profiling, hormone evaluation and cell proliferation observation revealed that disturbing ZmECR1 expression mainly affects pathways on hormone signal transduction, cell cycle progression and starch accumulation during kernel development. In addition, mutant in zmaxr1 (Auxin resistant 1), another RUB E1 subunit, also showed similar defects in kernel development. Double mutation of zmecr1 and zmaxr1 lead to empty pericarp kernel phenotype. RUBylation is a novel regulatory pathway affecting maize kernel development, majorly through its functions in modifying multiple cellular progresses.
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Claims 7
Claim · model-written summary
Smk501 encodes the RUBylation activating enzyme E1 subunit ZmECR1.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Smk501 encodes a RUBylation activating enzyme E1 subunit ZmECR1 (E1 C-TERMINAL RELATED 1) protein”
t0“maize”
Claim · model-written summary
The smk501 mutation produced fewer embryo and endosperm cells.
- Material
- smk501 mutatione0
- Organ / stage
- embryo and endosperme0
- Gene
- smk501e0
- Outcome
- cell numbere0
- Direction
- lesse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Destruction in RUBylation by smk501 mutation resulted in less embryo and endosperm cell number and smaller kernel size”
t0“maize”
Claim · model-written summary
The smk501 mutation produced smaller maize kernels.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Destruction in RUBylation by smk501 mutation resulted in less embryo and endosperm cell number and smaller kernel size”
t0“maize”
Claim · model-written summary
Disturbed ZmECR1 expression mainly affected hormone signal transduction, cell cycle progression, and starch accumulation pathways during kernel development.
- Organ / stage
- kernel developmente0
- Gene
- ZmECR1e0
- Outcome
- hormone signal transduction, cell cycle progression and starch accumulatione0
Not reportedTaxon
Verbatim
e0“disturbing ZmECR1 expression mainly affects pathways on hormone signal transduction, cell cycle progression and starch accumulation during kernel development”
Claim · model-written summary
A zmaxr1 mutant showed similar kernel development defects.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“mutant in zmaxr1 (Auxin resistant 1), another RUB E1 subunit, also showed similar defects in kernel development”
t0“maize”
Claim · model-written summary
The zmecr1 and zmaxr1 double mutation caused an empty pericarp kernel phenotype.
- Material
- Double mutation of zmecr1 and zmaxr1e0
- Gene
- zmecr1 and zmaxr1e0
- Outcome
- empty pericarp kernel phenotypee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Double mutation of zmecr1 and zmaxr1 lead to empty pericarp kernel phenotype”
t0“maize”
Claim · model-written summary
The authors identify RUBylation as a regulatory pathway for maize kernel development through effects on multiple cellular processes.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“RUBylation is a novel regulatory pathway affecting maize kernel development, majorly through its functions in modifying multiple cellular progresses”