Paper · 2026 · Crop Design

Long-term elevated daytime temperature disrupts reproductive development and 2-AP biosynthesis gene expression in aromatic rice ‘KDML105’

Phithak Inthima; Natchanon Meesa; Kumrop Ratanasut; et al.

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Open access · cc-by

Abstract

Global warming poses a critical threat to rice production, with elevated daytime temperatures during reproduction causing irreversible yield losses. The premium aromatic cultivar ‘Khao Dawk Mali 105’ (‘KDML105’) is widely cultivated in Thailand but is highly heat-sensitive. To assess its vulnerability, plants were grown under controlled environments at 32, 36, and 40 °C from panicle initiation to maturity. Daytime warming ≥36 °C delayed flowering, shortened culm and panicle length, and inhibited uppermost internode elongation, resulting in incomplete panicle exsertion. Anther development was strongly affected, with reduced size, collapse of structural integrity, and poor pollen release at 36 °C, progressing to deformation and complete sterility at 40 °C. Spikelet fertility declined sharply with rising temperature, culminating in 100% sterility at 40 °C, confirming the narrow thermal threshold of ‘KDML105’. At the molecular level, elevated temperature downregulated proline–GABA pathway genes ( OsP5cs , OsProDH , OsOat , Osbadh2 ), suppressed biosynthesis of 2-acetyl-1-pyrroline (2AP), and increased volatilization losses, thereby diminishing aroma. Together, these results show that long-term elevated daytime temperature simultaneously compromises reproductive success and fragrance quality. Panicle traits and anther morphology serve as sensitive indicators for screening heat tolerance, while targeted breeding and adaptive agronomic strategies are essential to protect yield and grain quality in aromatic rice under future climate scenarios. • Daytime temperature ≥36 °C) during reproduction caused complete sterility in ‘KDML105’ rice. • High temperature reduced anther size, impaired dehiscence, and lowered pollen viability. • Culm elongation and panicle exsertion were suppressed, restricting pollination efficiency. • Elevated temperature downregulated proline–GABA pathway genes, limiting 2AP biosynthesis. • Long-term daytime warming threatens both yield and aroma quality in aromatic rice.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 11

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Warming at least 36 °C delayed flowering.

Material
KDML105t0
Treatment
Daytime warming ≥36 °Ce0
Outcome
floweringe0
Direction
delayede0

Not reportedComparator

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

“Daytime warming ≥36 °C delayed flowering”

✓ located verbatim in the source · abstract, characters 410–450 · show in text ↑
e1

“Daytime warming ≥36 °C delayed flowering”

✓ located verbatim in the source · abstract, characters 410–450 · show in text ↑
t0

“rice ‘KDML105’”

✓ located verbatim in the source · title, characters 123–137 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Warming shortened culm and panicle length.

Treatment
Daytime warming ≥36 °Ce0
Outcome
culm and panicle lengthe0
Direction
shortenede0

Not reportedComparator

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

“Daytime warming ≥36 °C delayed flowering, shortened culm and panicle length”

✓ located verbatim in the source · abstract, characters 410–485 · show in text ↑
e1

“Daytime warming ≥36 °C delayed flowering, shortened culm and panicle length”

✓ located verbatim in the source · abstract, characters 410–485 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Warming inhibited uppermost internode elongation and led to incomplete panicle exsertion.

Treatment
Daytime warming ≥36 °Ce1
Outcome
uppermost internode elongatione0
Direction
inhibitede0

Not reportedComparator

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

“inhibited uppermost internode elongation, resulting in incomplete panicle exsertion”

✓ located verbatim in the source · abstract, characters 491–574 · show in text ↑
e1

“Daytime warming ≥36 °C delayed flowering”

✓ located verbatim in the source · abstract, characters 410–450 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

At 36 °C, anthers were smaller and structurally collapsed with poor pollen release.

Organ / stage
Anthere1
Treatment
36 °Ce0
Outcome
size, collapse of structural integrity, and poor pollen releasee0
Direction
reducede0

Not reportedComparator

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

“reduced size, collapse of structural integrity, and poor pollen release at 36 °C”

✓ located verbatim in the source · abstract, characters 623–703 · show in text ↑
e1

“Anther development was strongly affected”

✓ located verbatim in the source · abstract, characters 576–616 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

At 40 °C, anthers became deformed and completely sterile.

Treatment
40 °Ce0
Outcome
deformation and complete sterilitye0

Not reportedComparatorDirection

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

“progressing to deformation and complete sterility at 40 °C”

✓ located verbatim in the source · abstract, characters 705–763 · show in text ↑
e1

“progressing to deformation and complete sterility at 40 °C”

✓ located verbatim in the source · abstract, characters 705–763 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Spikelet fertility declined with rising temperature, reaching 100% sterility at 40 °C.

Treatment
rising temperaturee0
Outcome
Spikelet fertilitye0
Direction
declinede0
Value
100% sterilitye0

Not reportedComparator

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

“Spikelet fertility declined sharply with rising temperature, culminating in 100% sterility at 40 °C”

✓ located verbatim in the source · abstract, characters 765–864 · show in text ↑
e1

“Spikelet fertility declined sharply with rising temperature, culminating in 100% sterility at 40 °C”

✓ located verbatim in the source · abstract, characters 765–864 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Elevated temperature downregulated four proline–GABA pathway genes.

Treatment
elevated temperaturee0
Gene
OsP5cs , OsProDH , OsOat , Osbadh2e0
Outcome
proline–GABA pathway genese0
Direction
downregulatede0

Not reportedComparator

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

“elevated temperature downregulated proline–GABA pathway genes ( OsP5cs , OsProDH , OsOat , Osbadh2 )”

✓ located verbatim in the source · abstract, characters 944–1044 · show in text ↑
e1

“elevated temperature downregulated proline–GABA pathway genes ( OsP5cs , OsProDH , OsOat , Osbadh2 )”

✓ located verbatim in the source · abstract, characters 944–1044 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Elevated temperature suppressed 2AP biosynthesis.

Treatment
elevated temperaturee1
Outcome
biosynthesis of 2-acetyl-1-pyrroline (2AP)e0
Direction
suppressede0

Not reportedComparator

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

“suppressed biosynthesis of 2-acetyl-1-pyrroline (2AP)”

✓ located verbatim in the source · abstract, characters 1046–1099 · show in text ↑
e1

“elevated temperature downregulated proline–GABA pathway genes ( OsP5cs , OsProDH , OsOat , Osbadh2 )”

✓ located verbatim in the source · abstract, characters 944–1044 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Elevated temperature increased volatilization losses.

Treatment
elevated temperaturee1
Outcome
volatilization lossese0
Direction
increasede0

Not reportedComparator

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

“increased volatilization losses, thereby diminishing aroma”

✓ located verbatim in the source · abstract, characters 1105–1163 · show in text ↑
e1

“elevated temperature downregulated proline–GABA pathway genes ( OsP5cs , OsProDH , OsOat , Osbadh2 )”

✓ located verbatim in the source · abstract, characters 944–1044 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Authors identify panicle traits and anther morphology as indicators for heat tolerance screening.

Outcome
Panicle traits and anther morphologye0

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

“Panicle traits and anther morphology serve as sensitive indicators for screening heat tolerance”

✓ located verbatim in the source · abstract, characters 1309–1404 · show in text ↑
e1

“Panicle traits and anther morphology serve as sensitive indicators for screening heat tolerance”

✓ located verbatim in the source · abstract, characters 1309–1404 · show in text ↑
t0

“rice”

✓ located verbatim in the source · title, characters 123–127 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Authors recommend targeted breeding and adaptive agronomic strategies.

Outcome
targeted breeding and adaptive agronomic strategiese0

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

“targeted breeding and adaptive agronomic strategies are essential to protect yield and grain quality in aromatic rice”

✓ located verbatim in the source · abstract, characters 1412–1529 · show in text ↑
e1

“targeted breeding and adaptive agronomic strategies are essential to protect yield and grain quality in aromatic rice”

✓ located verbatim in the source · abstract, characters 1412–1529 · show in text ↑

heat · molecular: metabolites · Oryza sativa (rice)Results differ

43 papers: 13 positive, 30 negative or no effect

29 papers in the same direction as this paper · 13 in the opposite direction

  1. negative
  2. positive
  3. positive

39 papers more in this group

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heat · phenology · Oryza sativa (rice)Positive and negative results

6 papers: 1 positive, 5 negative or no effect

4 papers in the same direction as this paper · 1 in the opposite direction

  1. positive
  2. negative
  3. negative

2 papers more in this group

See all 6 papers →

heat · molecular: expression · Oryza sativa (rice)Positive and negative results

35 papers: 17 positive, 18 negative or no effect

17 papers in the same direction as this paper · 17 in the opposite direction

  1. no effect
  2. positive
  3. positive

31 papers more in this group

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heat · growth · Oryza sativa (rice)Positive and negative results

107 papers: 46 positive, 61 negative or no effect

60 papers in the same direction as this paper · 46 in the opposite direction

  1. negative
  2. positive
  3. positive

103 papers more in this group

See all 107 papers →

heat · yield · Oryza sativa (rice)Positive and negative results

195 papers: 36 positive, 159 negative or no effect

158 papers in the same direction as this paper · 36 in the opposite direction

  1. negative
  2. negative
  3. negative

191 papers more in this group

See all 195 papers →

Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.