Paper · 2026 · Ecological Research

Physiological and Nutritional Mechanisms Underlying the Crimson Coloration of Boxwood on Ultramafic Soils of Sugashima Island, Japan

Takafumi Mizuno; Yuki Minatani; Mayuko Hara

10
claims
18
verbatim quotes
0
citations
2
research areas
✓
Open access

Abstract

ABSTRACT The evergreen shrub Buxus microphylla (Buxaceae; Japanese boxwood) forms distinctive ultramafic vegetation in Japan, with populations on Sugashima Island exhibiting unusually vivid crimson leaves in winter. To clarify the physiological and nutritional mechanisms underlying this coloration, we combined field monitoring with laboratory experiments to reproduce leaf reddening under controlled conditions. Field observations revealed that leaves of B. microphylla growing on ultramafic soils turned bright red each winter, whereas those from non‐ultramafic sites remained mostly green, except for slight reddening at the Mie University site. The crimson coloration coincided with a seasonal decline in leaf nitrogen (N) and chlorophyll content, suggesting that nutrient limitation enhances pigment imbalance. In laboratory assays, light exposure and sucrose addition strongly promoted chlorophyll degradation, increasing the carotenoid‐to‐chlorophyll ratio (Car/Chl) by more than 20%. Under dark conditions, both sucrose and N enhanced pigment retention. The Car/Chl ratio consistently ranked as water‐only > N‐only > multi‐nutrient media, indicating that nutrient sufficiency counteracts sugar‐induced photoinhibition and chlorophyll breakdown. These results demonstrate that leaf reddening in boxwood is not due to anthocyanin synthesis but rather to a relative enrichment of carotenoids following chlorophyll loss under photoinhibitory conditions. The combination of high irradiance, low temperature, and N deficiency on ultramafic soils promotes this imbalance, producing the intense “crimson boxwood” phenotype. Our findings highlight how nutrient limitation and photoprotection interact to shape adaptive pigmentation in plants growing on ultramafic soils.

Abstract from the paper’s OpenAlex record, shown for indexing and research. Copyright remains with the authors or the publisher. 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 10

Associationobservedoriginal resultmeasured

Claim · model-written summary

B. microphylla leaves on ultramafic soils turned bright red each winter, whereas leaves from non-ultramafic sites remained mostly green.

Taxon
B. microphyllaBuxus microphyllae0
Outcome
turned bright red each wintere0
Comparator
non‐ultramafic sitese0
Setting
ultramafic soilse0

Not reportedDirection

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

Buxus microphylla
Buxus microphylla · Taxon · picture from its Wikimedia Commons categoryTesstree · CC0 · Wikimedia Commons ↗

Verbatim

e0

“leaves of B. microphylla growing on ultramafic soils turned bright red each winter, whereas those from non‐ultramafic sites remained mostly green”

✓ located verbatim in the source · abstract, characters 447–592 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Crimson coloration coincided with a seasonal decline in leaf nitrogen.

Taxon
Boxwoodt0
Organ / stage
leafe0
Outcome
leaf nitrogen (N)e0
Direction
declinee0
Qualifier
crimson coloratione0

Verbatim

e0

“The crimson coloration coincided with a seasonal decline in leaf nitrogen (N) and chlorophyll content”

✓ located verbatim in the source · abstract, characters 650–751 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Crimson coloration coincided with a seasonal decline in chlorophyll content.

Taxon
Boxwoodt0
Organ / stage
leafe0
Outcome
chlorophyll contente0
Direction
declinee0
Qualifier
crimson coloratione0

Verbatim

e0

“The crimson coloration coincided with a seasonal decline in leaf nitrogen (N) and chlorophyll content”

✓ located verbatim in the source · abstract, characters 650–751 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Light exposure and sucrose addition promoted chlorophyll degradation.

Taxon
Boxwoodt0
Treatment
light exposure and sucrose additione0
Outcome
chlorophyll degradatione0
Direction
promotede0

Not reportedComparator

Verbatim

e0

“light exposure and sucrose addition strongly promoted chlorophyll degradation, increasing the carotenoid‐to‐chlorophyll ratio (Car/Chl) by more than 20%”

✓ located verbatim in the source · abstract, characters 839–991 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Light exposure and sucrose addition increased the Car/Chl ratio by more than 20%.

Taxon
Boxwoodt0
Treatment
light exposure and sucrose additione0
Outcome
carotenoid‐to‐chlorophyll ratio (Car/Chl)e0
Direction
increasinge0
Value
more than 20%e0

Not reportedComparator

Verbatim

e0

“light exposure and sucrose addition strongly promoted chlorophyll degradation, increasing the carotenoid‐to‐chlorophyll ratio (Car/Chl) by more than 20%”

✓ located verbatim in the source · abstract, characters 839–991 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Under darkness, sucrose enhanced pigment retention.

Taxon
Boxwoodt0
Treatment
sucrosee0
Outcome
pigment retentione0
Direction
enhancede0
Qualifier
dark conditionse0

Not reportedComparator

Verbatim

e0

“Under dark conditions, both sucrose and N enhanced pigment retention”

✓ located verbatim in the source · abstract, characters 993–1061 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Under darkness, nitrogen enhanced pigment retention.

Taxon
Boxwoodt0
Treatment
Ne0
Outcome
pigment retentione0
Direction
enhancede0
Qualifier
dark conditionse0

Not reportedComparator

Verbatim

e0

“Under dark conditions, both sucrose and N enhanced pigment retention”

✓ located verbatim in the source · abstract, characters 993–1061 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

The Car/Chl ratio ranked highest in water-only medium, followed by N-only medium, then multi-nutrient medium.

Taxon
Boxwoodt0
Outcome
Car/Chl ratioe0
Direction
water‐only > N‐only > multi‐nutrient mediae0

Verbatim

e0

“The Car/Chl ratio consistently ranked as water‐only > N‐only > multi‐nutrient media”

✓ located verbatim in the source · abstract, characters 1063–1146 · show in text ↑
t0

“Boxwood”

✓ located verbatim in the source · title, characters 78–85 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

The authors attribute red boxwood leaves to relative carotenoid enrichment after chlorophyll loss rather than anthocyanin synthesis.

Taxon
boxwoode0
Outcome
leaf reddeninge0
Qualifier
relative enrichment of carotenoids following chlorophyll loss under photoinhibitory conditionse0

Verbatim

e0

“leaf reddening in boxwood is not due to anthocyanin synthesis but rather to a relative enrichment of carotenoids following chlorophyll loss under photoinhibitory conditions”

✓ located verbatim in the source · abstract, characters 1285–1457 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

The authors infer that high irradiance, low temperature and nitrogen deficiency on ultramafic soils promote the crimson phenotype.

Taxon
boxwoode0
Outcome
“crimson boxwood” phenotypee0
Qualifier
high irradiance, low temperature, and N deficiency on ultramafic soilse0

Verbatim

e0

“The combination of high irradiance, low temperature, and N deficiency on ultramafic soils promotes this imbalance, producing the intense “crimson boxwood” phenotype”

✓ located verbatim in the source · abstract, characters 1459–1623 · show in text ↑