Paper · 2026 · Journal of Genetic Engineering and Biotechnology

Morpho-molecular diversity of improved Indonesia’s rice varieties based on grain morphology and three types of DNA-based markers

Reflinur Reflinur; Adin Heriyan Nugroho; Zidni Muflikhati; et al.

19
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Open access · cc-by-nc-nd

Abstract

Genetic improvement of rice (Oryza sativa L.) requires a clear understanding of both phenotypic and molecular diversity. This study examined 82 improved Indonesian rice varieties using grain morphological traits and three DNA-based marker systems: Simple Sequence Repeat (SSR), Subspecies-Specific Sequence-Tagged Site (SS-STS), and Start Codon Targeted (SCoT). Phenotypic evaluation revealed significant variation in grain length, width, and thickness. Principal component analysis condensed four traits into two axes explaining 94.4% of variance, separating genotypes into three clusters largely corresponding to indica and japonica types. Across 32 markers, 157 alleles were detected, with average gene diversity (He) values of 0.30 (SSR), 0.29 (SS-STS), and 0.22 (SCoT). Polymorphic information content (PIC) was generally low. SS-STS markers revealed fixation of indica-type alleles, consistent with major varieties IR64 and Ciherang, while rare alleles were scarce (1.1-1.5 per locus). Although SCoT markers generated 8.5 bands per locus, heterozygosity remained low (0.005-0.010). Analysis of molecular variance (AMOVA) showed that 90-91% of the genetic variation was distributed within subpopulations, while only 5-8% was explained by differences among subpopulations, indicating weak population structure (FST = 0.055 for SSR; 0.084 for SS-STS). STRUCTURE analysis inferred different subpopulation numbers for each marker type (K = 7 for SS-STS, K = 2 for SSR, K = 4 for SCoT). Phylogenetic clustering separated the genotypes into two to three subpopulations, which partly corresponded to grain type and inferred subspecies background. These results reveal limited molecular diversity despite observable phenotypic variation, suggesting genetic narrowing associated with the repeated use of closely related parental lines during breeding. Broader germplasm, including landraces and wild relatives, is essential for future rice breeding.

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Claims 19

Observationobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

The 82 improved Indonesian rice varieties showed significant variation in grain length, width and thickness.

Outcome
variation in grain length, width, and thicknesse0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“Phenotypic evaluation revealed significant variation in grain length, width, and thickness”

✓ located verbatim in the source · abstract, characters 362–452 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Principal component analysis condensed the four grain traits into two axes explaining 94.4% of variance.

Outcome
variancee0
Value
94.4%e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“Principal component analysis condensed four traits into two axes explaining 94.4% of variance”

✓ located verbatim in the source · abstract, characters 454–547 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Principal component analysis separated the genotypes into three clusters largely corresponding to indica and japonica types.

Outcome
three clusters largely corresponding to indica and japonica typese0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“separating genotypes into three clusters largely corresponding to indica and japonica types”

✓ located verbatim in the source · abstract, characters 549–640 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Across 32 markers, 157 alleles were detected in the rice varieties.

Outcome
alleles were detectede0
Value
157 allelese0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“Across 32 markers, 157 alleles were detected”

✓ located verbatim in the source · abstract, characters 642–686 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Average gene diversity (He) was 0.30 with SSR markers.

Outcome
gene diversity (He)e0
Value
0.30 (SSR)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“average gene diversity (He) values of 0.30 (SSR), 0.29 (SS-STS), and 0.22 (SCoT)”

✓ located verbatim in the source · abstract, characters 693–773 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Average gene diversity (He) was 0.29 with SS-STS markers.

Outcome
gene diversity (He)e0
Value
0.29 (SS-STS)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“average gene diversity (He) values of 0.30 (SSR), 0.29 (SS-STS), and 0.22 (SCoT)”

✓ located verbatim in the source · abstract, characters 693–773 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Average gene diversity (He) was 0.22 with SCoT markers.

Outcome
gene diversity (He)e0
Value
0.22 (SCoT)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“average gene diversity (He) values of 0.30 (SSR), 0.29 (SS-STS), and 0.22 (SCoT)”

✓ located verbatim in the source · abstract, characters 693–773 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Polymorphic information content was generally low.

Outcome
Polymorphic information content (PIC)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“Polymorphic information content (PIC) was generally low”

✓ located verbatim in the source · abstract, characters 775–830 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

SS-STS markers showed fixation of indica-type alleles, consistent with the major varieties IR64 and Ciherang.

Material
IR64 and Ciherange0
Outcome
fixation of indica-type allelese0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“SS-STS markers revealed fixation of indica-type alleles, consistent with major varieties IR64 and Ciherang”

✓ located verbatim in the source · abstract, characters 832–938 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Rare alleles were scarce, at 1.1-1.5 per locus.

Outcome
rare allelese0
Value
1.1-1.5 per locuse0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“rare alleles were scarce (1.1-1.5 per locus)”

✓ located verbatim in the source · abstract, characters 946–990 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

SCoT markers generated 8.5 bands per locus.

Outcome
bands per locuse0
Value
8.5e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“SCoT markers generated 8.5 bands per locus”

✓ located verbatim in the source · abstract, characters 1001–1043 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Heterozygosity remained low, at 0.005-0.010.

Outcome
heterozygositye0
Value
0.005-0.010e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“heterozygosity remained low (0.005-0.010)”

✓ located verbatim in the source · abstract, characters 1045–1086 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

AMOVA showed 90-91% of the genetic variation was distributed within subpopulations.

Outcome
genetic variation was distributed within subpopulationse0
Value
90-91%e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“90-91% of the genetic variation was distributed within subpopulations”

✓ located verbatim in the source · abstract, characters 1139–1208 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Only 5-8% of the genetic variation was explained by differences among subpopulations.

Outcome
explained by differences among subpopulationse0
Value
5-8%e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“only 5-8% was explained by differences among subpopulations”

✓ located verbatim in the source · abstract, characters 1216–1275 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Population structure was weak, with FST of 0.055 for SSR and 0.084 for SS-STS.

Outcome
population structuree0
Value
FST = 0.055 for SSR; 0.084 for SS-STSe0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“indicating weak population structure (FST = 0.055 for SSR; 0.084 for SS-STS)”

✓ located verbatim in the source · abstract, characters 1277–1353 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

STRUCTURE analysis inferred different numbers of subpopulations for each marker type (K = 7 for SS-STS, K = 2 for SSR, K = 4 for SCoT).

Outcome
subpopulation numberse0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“STRUCTURE analysis inferred different subpopulation numbers for each marker type (K = 7 for SS-STS, K = 2 for SSR, K = 4 for SCoT)”

✓ located verbatim in the source · abstract, characters 1355–1485 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Phylogenetic clustering separated the genotypes into two to three subpopulations, partly corresponding to grain type and inferred subspecies background.

Outcome
Phylogenetic clustering separated the genotypes into two to three subpopulationse0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“Phylogenetic clustering separated the genotypes into two to three subpopulations, which partly corresponded to grain type and inferred subspecies background”

✓ located verbatim in the source · abstract, characters 1487–1643 · show in text ↑
Observationauthor interpretationoriginal resultmeasured

Claim · model-written summary

The authors conclude molecular diversity is limited despite phenotypic variation, suggesting genetic narrowing from repeated use of closely related parental lines in breeding.

Outcome
limited molecular diversitye0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

t0

“rice varieties”

✓ located verbatim in the source · title, characters 51–65 · show in text ↑
e0

“These results reveal limited molecular diversity despite observable phenotypic variation, suggesting genetic narrowing associated with the repeated use of closely related parental lines during breeding”

✓ located verbatim in the source · abstract, characters 1645–1846 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Broader germplasm, including landraces and wild relatives, is essential for future rice breeding.

Outcome
Broader germplasm, including landraces and wild relativese0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“Broader germplasm, including landraces and wild relatives, is essential for future rice breeding”

✓ located verbatim in the source · abstract, characters 1848–1944 · show in text ↑