Abstract
PREMISE OF THE STUDY: Mixotrophy is a strategy whereby plants acquire carbon both through photosynthesis and heterotrophic exploitation of mycorrhizal fungi. In Euro-American Pyroleae species studied hitherto, heterotrophy levels vary according to species, sites of study, and possibly light conditions. We investigated mycorrhizal association and mixotrophy in the Asiatic forest species Pyrola japonica, and their plasticity under different light conditions. METHODS: Pyrola japonica was sampled bimonthly in sunny and shaded conditions from a deciduous broadleaf forest. We microscopically assessed the rate of fungal colonization and sequenced the ITS to identify the mycorrhizal fungi. We measured (13)C and (15)N isotopic abundances in P. japonica as compared with neighboring autotrophic and mycoheterotrophic plants, to evaluate P. japonica's heterotrophy level. KEY RESULTS: Pyrola japonica formed arbutoid mycorrhizas devoid of fungal mantles, with intracellular hyphal coils and a Hartig net. It tended to be more colonized by mycorrhizal fungi in spring and summer. Most associated fungi belonged to ectomycorrhizal taxa, and 84% of identified fungi were Russula spp. Rate of mycorrhizal colonization and Russula frequency tended to be higher in shaded conditions. Both δ(13)C and δ(15)N values of P. japonica were significantly higher in autotrophic plants, showing that about half of the carbon on average was received from mycorrhizal fungi. Both isotopic values negatively correlated with light availability, suggesting higher heterotrophy levels in shaded conditions. CONCLUSIONS: The mixotrophic P. japonica undergoes changes in mycorrhizal symbionts and carbon nutrition according to light availability. Our results suggest that during Pyroleae evolution, a tendency to increased heterotrophy emerged in the Pyrola/Orthilia clade.
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Claims 11
Claim · model-written summary
Pyrola japonica formed arbutoid mycorrhizas without mantles, with hyphal coils and a Hartig net.
- Taxon
- Pyrola japonica
Pyrola japonicae0
- Interactor
- fungale0symbiont
- Outcome
- arbutoid mycorrhizas devoid of fungal mantles, with intracellular hyphal coils and a Hartig nete0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Pyrola japonica formed arbutoid mycorrhizas devoid of fungal mantles, with intracellular hyphal coils and a Hartig net”
Claim · model-written summary
Fungal colonization tended to be greater in spring and summer.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Interactor
- mycorrhizal fungie0symbiont
- Outcome
- colonizede0
- Direction
- moree0
- Setting
- spring and summere0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“It tended to be more colonized by mycorrhizal fungi in spring and summer”
t0“Pyrola japonica”
Claim · model-written summary
Most associated fungi were ectomycorrhizal taxa.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Interactor
- ectomycorrhizal taxae0symbiont
- Outcome
- associated fungi belonged to ectomycorrhizal taxae0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Most associated fungi belonged to ectomycorrhizal taxa”
t0“Pyrola japonica”
Claim · model-written summary
Russula species accounted for 84% of identified fungi.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Outcome
- identified fungi were Russula spp.e0
- Value
- 84%e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“84% of identified fungi were Russula spp.”
t0“Pyrola japonica”
Claim · model-written summary
Mycorrhizal colonization tended to be higher in shade.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Outcome
- Rate of mycorrhizal colonizatione0
- Direction
- highere0
- Setting
- shaded conditionse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Rate of mycorrhizal colonization and Russula frequency tended to be higher in shaded conditions”
t0“Pyrola japonica”
Claim · model-written summary
Russula frequency tended to be higher in shade.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Outcome
- Russula frequencye0
- Direction
- highere0
- Setting
- shaded conditionse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Rate of mycorrhizal colonization and Russula frequency tended to be higher in shaded conditions”
t0“Pyrola japonica”
Claim · model-written summary
Pyrola japonica δ13C and δ15N values were reported significantly higher than autotrophic plants.
- Taxon
- P. japonica
Pyrola japonicae0
- Outcome
- δ(13)C and δ(15)N valuese0
- Direction
- highere0
- Comparator
- autotrophic plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Both δ(13)C and δ(15)N values of P. japonica were significantly higher in autotrophic plants”
Claim · model-written summary
About half the carbon was inferred to come from mycorrhizal fungi.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Interactor
- mycorrhizal fungie0symbiont
- Outcome
- carbon on average was received from mycorrhizal fungie0
- Value
- about halfe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“about half of the carbon on average was received from mycorrhizal fungi”
t0“Pyrola japonica”
Claim · model-written summary
Both isotope values correlated negatively with light availability.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Outcome
- isotopic valuese0
- Direction
- negatively correlatede0
- Setting
- light availabilitye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Both isotopic values negatively correlated with light availability”
t0“Pyrola japonica”
Claim · model-written summary
The authors inferred higher heterotrophy under shaded conditions.
- Taxon
- Pyrola japonica
Pyrola japonicat0
- Outcome
- heterotrophy levelse0
- Direction
- highere0
- Setting
- shaded conditionse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“suggesting higher heterotrophy levels in shaded conditions”
t0“Pyrola japonica”
Claim · model-written summary
The authors suggested increased heterotrophy evolved in the Pyrola/Orthilia clade.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e0“during Pyroleae evolution, a tendency to increased heterotrophy emerged in the Pyrola/Orthilia clade”