Abstract
Background and Aims: The cuticular waxes sealing plant surfaces against excessive water loss are complex mixtures of very-long-chain aliphatics, with compositions that vary widely between plant species. To help fill the gap in our knowledge about waxes of non-flowering plant taxa, and thus about the cuticle of ancestral land plants, this study provides comprehensive analyses of waxes on temperate fern species from five different families. Methods: The wax mixtures on fronds of Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris were analysed using gas chromatography-mass spectrometry for identification, and gas chromatography-flame ionization detection for quantification. Key Results: The wax mixtures from all five fern species contained large amounts of C36-C54 alkyl esters, with species-specific homologue distributions. They were accompanied by minor amounts of fatty acids, primary alcohols, aldehydes and/or alkanes, whose chain length profiles also varied widely between species. In the frond wax of G. dryopteris, C27-C33 secondary alcohols and C27-C35 ketones with functional groups exclusively on even-numbered carbons (C-10 to C-16) were identified; these are characteristic structures similar to secondary alcohols and ketones in moss, gymnosperm and basal angiosperm waxes. The ferns had total wax amounts varying from 3.9 μg cm-2 on P. glycyrrhiza to 16.9 μg cm-2 on G. dryopteris, thus spanning a range comparable with that on leaves of flowering plants. Conclusions: The characteristic compound class compositions indicate that all five fern species contain the full complement of wax biosynthesis enzymes previously described for the angiosperm arabidopsis. Based on the isomer profiles, we predict that each fern species, in contrast to arabidopsis, has multiple ester synthase enzymes, each with unique substrate specificities.
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Claims 7
Claim · model-written summary
All five fern species had abundant C36-C54 alkyl esters with species-specific homologue distributions.
- Taxon
- Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris
Polystichum munitum
Pteridium aquilinumGymnocarpium dryopteris
Polypodium glycyrrhiza
Cryptogramma crispat0 - Outcome
- large amounts of C36-C54 alkyl esterse0
- Qualifier
- species-specific homologue distributionse0
Not reportedOrgan / stage
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
_001.jpg?width=500)


Verbatim
e0“The wax mixtures from all five fern species contained large amounts of C36-C54 alkyl esters, with species-specific homologue distributions.”
t0“Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris”
Claim · model-written summary
Minor fatty acids, primary alcohols, aldehydes and/or alkanes had chain-length profiles that varied widely between species.
- Taxon
- Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris
Polystichum munitum
Pteridium aquilinumGymnocarpium dryopteris
Polypodium glycyrrhiza
Cryptogramma crispat0 - Outcome
- fatty acids, primary alcohols, aldehydes and/or alkanese0
- Qualifier
- chain length profiles also varied widely between speciese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
_001.jpg?width=500)


Verbatim
e0“They were accompanied by minor amounts of fatty acids, primary alcohols, aldehydes and/or alkanes, whose chain length profiles also varied widely between species.”
t0“Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris”
Claim · model-written summary
G. dryopteris frond wax contained C27-C33 secondary alcohols and C27-C35 ketones with functional groups exclusively on even-numbered carbons.
- Taxon
- G. dryopteris
Gymnocarpium dryopterise0
- Organ / stage
- fronde0
- Outcome
- C27-C33 secondary alcohols and C27-C35 ketonese0
- Qualifier
- functional groups exclusively on even-numbered carbons (C-10 to C-16)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“In the frond wax of G. dryopteris, C27-C33 secondary alcohols and C27-C35 ketones with functional groups exclusively on even-numbered carbons (C-10 to C-16) were identified”
t0“Gymnocarpium dryopteris”
Claim · model-written summary
Frond wax coverage was 3.9 μg cm-2 on P. glycyrrhiza.
- Taxon
- P. glycyrrhiza
Polypodium glycyrrhizae0 - Organ / stage
- frondst0
- Outcome
- Coveraget0
- Value
- 3.9 μg cm-2e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“3.9 μg cm-2 on P. glycyrrhiza”
t0“Coverage and composition of cuticular waxes on the fronds”
Claim · model-written summary
Frond wax coverage was 16.9 μg cm-2 on G. dryopteris.
- Taxon
- G. dryopteris
Gymnocarpium dryopterise0
- Organ / stage
- frondst0
- Outcome
- Coveraget0
- Value
- 16.9 μg cm-2e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“16.9 μg cm-2 on G. dryopteris”
t0“Coverage and composition of cuticular waxes on the fronds”
Claim · model-written summary
The five ferns appear to have the full complement of wax biosynthesis enzymes described for arabidopsis.
- Taxon
- Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris
Polystichum munitum
Pteridium aquilinumGymnocarpium dryopteris
Polypodium glycyrrhiza
Cryptogramma crispat0 - Gene
- wax biosynthesis enzymese0
- Outcome
- full complement of wax biosynthesis enzymese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
_001.jpg?width=500)


Verbatim
e0“all five fern species contain the full complement of wax biosynthesis enzymes previously described for the angiosperm arabidopsis”
t0“Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris”
Claim · model-written summary
Each fern species is predicted to have multiple ester synthase enzymes with unique substrate specificities.
- Taxon
- Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris
Polystichum munitum
Pteridium aquilinumGymnocarpium dryopteris
Polypodium glycyrrhiza
Cryptogramma crispat0 - Gene
- ester synthase enzymese0
- Outcome
- multiple ester synthase enzymese0
- Qualifier
- unique substrate specificitiese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
_001.jpg?width=500)


Verbatim
e0“we predict that each fern species, in contrast to arabidopsis, has multiple ester synthase enzymes, each with unique substrate specificities”
t0“Pteridium aquilinum, Cryptogramma crispa, Polypodium glycyrrhiza, Polystichum munitum and Gymnocarpium dryopteris”