Abstract
MADS-box transcription factors (TFs) are involved in multiple plant development processes and are most known during the reproductive transition and floral organ development. Very few genes have been characterized in the genome of Humulus lupulus L. (Cannabaceae), an important crop for the pharmaceutical and beverage industries. The MADS-box family has not been studied in this species yet. We identified 65 MADS-box genes in the hop genome, of which 29 encode type-II TFs (27 of subgroup MIKCC and 2 MIKC*) and 36 type-I proteins (26 α, 9 β, and 1 γ). Type-II MADS-box genes evolved more complex architectures than type-I genes. Interestingly, we did not find FLOWERING LOCUS C (FLC) homologs, a transcription factor that acts as a floral repressor and is negatively regulated by cold. This result provides a molecular explanation for a previous work showing that vernalization is not a requirement for hop flowering, which has implications for its cultivation in the tropics. Analysis of gene ontology and expression profiling revealed genes potentially involved in the development of male and female floral structures based on the differential expression of ABC homeotic genes in each whorl of the flower. We identified a gene exclusively expressed in lupulin glands, suggesting a role in specialized metabolism in these structures. In toto, this work contributes to understanding the evolutionary history of MADS-box genes in hop, and provides perspectives on functional genetic studies, biotechnology, and crop breeding.
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Claims 8
Claim · model-written summary
The hop genome contained 65 MADS-box genes.
- Taxon
- Humulus lupulus L.
Humulus lupuluse1 - Gene
- MADS-box genese0
- Outcome
- MADS-box genese0
- Value
- 65e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“We identified 65 MADS-box genes in the hop genome”
e1“Humulus lupulus L.”
Claim · model-written summary
Of these genes, 29 encode type-II transcription factors, comprising 27 MIKCC and two MIKC*.
- Taxon
- Humulus lupulus L.
Humulus lupuluse1 - Outcome
- type-II TFse0
- Value
- 29e0
- Qualifier
- 27 of subgroup MIKCC and 2 MIKC*e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“29 encode type-II TFs (27 of subgroup MIKCC and 2 MIKC*)”
e1“Humulus lupulus L.”
Claim · model-written summary
Another 36 encode type-I proteins, comprising 26 alpha, nine beta and one gamma.
- Taxon
- Humulus lupulus L.
Humulus lupuluse1 - Outcome
- type-I proteinse0
- Value
- 36e0
- Qualifier
- 26 α, 9 β, and 1 γe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“36 type-I proteins (26 α, 9 β, and 1 γ)”
e1“Humulus lupulus L.”
Claim · model-written summary
Type-II MADS-box genes evolved more complex architectures than type-I genes.
- Taxon
- Humulus lupulus L.
Humulus lupuluse1 - Gene
- MADS-box genese0
- Outcome
- architecturese0
- Direction
- more complexe0
- Comparator
- type-I genese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Type-II MADS-box genes evolved more complex architectures than type-I genes.”
e1“Humulus lupulus L.”
Claim · model-written summary
No FLOWERING LOCUS C homologs were found in hop.
Verbatim
e0“we did not find FLOWERING LOCUS C (FLC) homologs”
e1“hop genome”
Claim · model-written summary
The absence of FLC homologs was proposed to explain lack of a vernalization requirement.
Verbatim
e0“This result provides a molecular explanation for a previous work showing that vernalization is not a requirement for hop flowering”
e1“FLOWERING LOCUS C (FLC) homologs”
Claim · model-written summary
Expression data indicated candidate genes for male and female floral structure development.
- Taxon
- Humulus lupulus L.
Humulus lupuluse1 - Gene
- ABC homeotic genese0
- Outcome
- development of male and female floral structurese0
- Qualifier
- differential expressione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“genes potentially involved in the development of male and female floral structures based on the differential expression of ABC homeotic genes in each whorl of the flower.”
e1“Humulus lupulus L.”
Claim · model-written summary
One gene was expressed exclusively in lupulin glands, suggesting specialized metabolism.
- Taxon
- Humulus lupulus L.
Humulus lupuluse1 - Gene
- a genee0
- Outcome
- exclusively expressed in lupulin glandse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“We identified a gene exclusively expressed in lupulin glands, suggesting a role in specialized metabolism in these structures.”
e1“Humulus lupulus L.”