Abstract
The angiosperm calyces display considerable diversity and have adaptive functions. However, the evolutionary trajectories and underlying mechanisms of calyx morphological diversity remain unclear. In this study, ancestral state reconstruction revealed that the abscised calyx was ancestral; however, most extant angiosperms exhibited persistent calyces showing notable variation in size. Remarkably, the Solanaceae family may represent a miniature reflecting the calyx diversity of angiosperms. Distinct from Solanum and Capsicum, Physalis fruits featured a morphological novelty known as inflated calyx syndrome (ICS). To reveal the molecular repatterning events underlying ICS formation, we conducted time-course transcriptomic comparisons on developing calyces of ICS species (Physalis floridana) and non-ICS species (Capsicum annuum and two Solanum species), and detected that variations in heterometric expression and alternative splicing were predominant across these species. Moreover, two Physalis-calyx highly expressed genes respectively encoding PHYSALIS ORGAN SIZE 4 (POS4) and POS5 were knocked down and out using virus-induced gene silencing and CRISPR/Cas9 technologies, and the resulting genetically modified P. floridana plant lines displayed a significant reduction in ICS size. Furthermore, when compared with Solanum and Capsicum, heterotopically expressed genes in Physalis calyx relative to berry were mainly enriched for functions in photosynthesis and responses to stimuli, thereby supporting the hypothesis that the inflated fruiting calyx may have partitioned and exapted functions originally associated with berry. This work elucidates the calyx evolutionary pattern of angiosperms as well as transcriptomic repatterning mechanisms that may govern both developmental and functional evolution of fruiting calyx inflation within Solanaceae, thereby providing insights into plant morphological evolution.
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Claims 8
Claim · model-written summary
Ancestral state reconstruction indicated that the abscised calyx was ancestral in angiosperms.
- Taxon
- angiospermsMagnoliopsidat0
- Organ / stage
- calyxt0
- Outcome
- abscised calyx was ancestrale0
Verbatim
t0“Evolution of calyx diversity in angiosperms”
e0“ancestral state reconstruction revealed that the abscised calyx was ancestral”
Claim · model-written summary
Most extant angiosperms have persistent calyces with notable variation in size.
- Taxon
- angiospermsMagnoliopsidae0
- Outcome
- persistent calycese0
- Qualifier
- notable variation in sizee0
Verbatim
e0“most extant angiosperms exhibited persistent calyces showing notable variation in size”
Claim · model-written summary
The Solanaceae family may represent a miniature of angiosperm calyx diversity.
- Taxon
- SolanaceaeSolanaceaee0
- Outcome
- calyx diversitye0
Verbatim
e0“the Solanaceae family may represent a miniature reflecting the calyx diversity of angiosperms”
Claim · model-written summary
Physalis fruits have a morphological novelty, the inflated calyx syndrome (ICS), unlike Solanum and Capsicum.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Distinct from Solanum and Capsicum, Physalis fruits featured a morphological novelty known as inflated calyx syndrome (ICS)”
Claim · model-written summary
Time-course transcriptome comparison of developing calyces of Physalis floridana and non-ICS species found that heterometric expression and alternative splicing variations were predominant.
Verbatim
e0“we conducted time-course transcriptomic comparisons on developing calyces of ICS species (Physalis floridana) and non-ICS species (Capsicum annuum and two Solanum species), and detected that variations in heterometric expression and alternative splicing were predominant across these species”
Claim · model-written summary
Knockdown and knockout of POS4 and POS5 significantly reduced ICS size in P. floridana.
- Taxon
- P. floridana
Physalis pubescense1 - Treatment
- knocked down and out using virus-induced gene silencing and CRISPR/Cas9 technologiese1
- Gene
- PHYSALIS ORGAN SIZE 4 (POS4) and POS5e0
- Outcome
- ICS sizee1
- Direction
- reductione1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“PHYSALIS ORGAN SIZE 4 (POS4) and POS5”
e1“were knocked down and out using virus-induced gene silencing and CRISPR/Cas9 technologies, and the resulting genetically modified P. floridana plant lines displayed a significant reduction in ICS size”
Claim · model-written summary
Compared with Solanum and Capsicum, heterotopically expressed genes in Physalis calyx relative to berry were mainly enriched for photosynthesis and responses to stimuli.
- Organ / stage
- calyx relative to berrye0
- Outcome
- enriched for functions in photosynthesis and responses to stimulie0
- Comparator
- Solanum and Capsicume0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“when compared with Solanum and Capsicum, heterotopically expressed genes in Physalis calyx relative to berry were mainly enriched for functions in photosynthesis and responses to stimuli”
Claim · model-written summary
The inflated fruiting calyx may have partitioned and exapted functions originally associated with berry.
- Taxon
- SolanaceaeSolanaceaet0
- Organ / stage
- berrye0
- Outcome
- inflated fruiting calyxe0
Verbatim
t0“within Solanaceae”
e0“the inflated fruiting calyx may have partitioned and exapted functions originally associated with berry”