Abstract
Summary Multiple independent and overlapping pollen rejection pathways contribute to unilateral interspecific incompatibility ( UI ). In crosses between tomato species, pollen rejection usually occurs when the female parent is self‐incompatible ( SI ) and the male parent self‐compatible ( SC ) (the ‘ SI × SC rule’). Additional, as yet unknown, UI mechanisms are independent of self‐incompatibility and contribute to UI between SC species or populations. We identified a major quantitative trait locus on chromosome 10 ( ui10.1 ) which affects pollen‐side UI responses in crosses between cultivated tomato, Solanum lycopersicum , and Solanum pennellii LA 0716, both of which are SC and lack S ‐ RN ase, the pistil determinant of S ‐specificity in Solanaceae. Here we show that ui10.1 is a farnesyl pyrophosphate synthase gene ( FPS 2 ) expressed in pollen. Expression is about 18‐fold higher in pollen of S. pennellii than in S. lycopersicum . Pollen with the hypomorphic S. lycopersicum allele is selectively eliminated on pistils of the F 1 hybrid, leading to transmission ratio distortion in the F 2 progeny. CRISPR /Cas9‐generated knockout mutants ( fps2 ) in S. pennellii LA 0716 are self‐sterile due to pollen rejection, but mutant pollen is fully functional on pistils of S. lycopersicum . F 2 progeny of S. lycopersicum × S. pennellii ( fps2 ) show reversed transmission ratio distortion due to selective elimination of pollen bearing the knockout allele. Overexpression of FPS 2 in S. lycopersicum pollen rescues the pollen elimination phenotype. FPS 2 ‐based pollen selectivity does not involve S‐ RN ase and has not been previously linked to UI . Our results point to an entirely new mechanism of interspecific pollen rejection in plants.
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Claims 8
Claim · model-written summary
ui10.1 is the pollen-expressed FPS2 gene.
- Taxon
- Solanum lycopersicum
Solanum lycopersicume1 - Organ / stage
- pollene0
- Gene
- FPS 2e0
- Outcome
- expressed in pollene0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ui10.1 is a farnesyl pyrophosphate synthase gene ( FPS 2 ) expressed in pollen”
e1“cultivated tomato, Solanum lycopersicum , and Solanum pennellii LA 0716”
Claim · model-written summary
FPS2 expression was about 18-fold higher in S. pennellii pollen.
- Taxon
- S. pennellii
Solanum pennelliie0 - Organ / stage
- pollene0
- Gene
- FPS 2e1
- Outcome
- Expressione0
- Direction
- highere0
- Value
- 18‐folde0
- Comparator
- S. lycopersicume0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Expression is about 18‐fold higher in pollen of S. pennellii than in S. lycopersicum”
e1“Overexpression of FPS 2 in S. lycopersicum pollen”
Claim · model-written summary
The S. lycopersicum allele pollen was eliminated on F1 hybrid pistils, distorting F2 transmission.
- Taxon
- S. lycopersicum
Solanum lycopersicume0 - Organ / stage
- Pollene0
- Outcome
- selectively eliminatede0
- Qualifier
- transmission ratio distortion in the F 2 progenye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Pollen with the hypomorphic S. lycopersicum allele is selectively eliminated on pistils of the F 1 hybrid, leading to transmission ratio distortion in the F 2 progeny”
Claim · model-written summary
S. pennellii fps2 knockouts were self-sterile from pollen rejection.
- Taxon
- S. pennellii
Solanum pennelliie0 - Treatment
- CRISPR /Cas9‐generated knockoute0
- Gene
- fps2e0
- Outcome
- self‐sterile due to pollen rejectione0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“CRISPR /Cas9‐generated knockout mutants ( fps2 ) in S. pennellii LA 0716 are self‐sterile due to pollen rejection”
Claim · model-written summary
S. pennellii fps2 mutant pollen functioned on S. lycopersicum pistils.
- Taxon
- S. pennellii
Solanum pennelliie1 - Organ / stage
- mutant pollene0
- Outcome
- fully functionale0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“mutant pollen is fully functional on pistils of S. lycopersicum”
e1“in S. pennellii LA 0716”
Claim · model-written summary
F2 progeny from the fps2 cross showed reversed transmission distortion.
- Taxon
- S. lycopersicum
Solanum lycopersicume0 - Gene
- fps2e0
- Outcome
- reversed transmission ratio distortione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“F 2 progeny of S. lycopersicum × S. pennellii ( fps2 ) show reversed transmission ratio distortion due to selective elimination of pollen bearing the knockout allele”
Claim · model-written summary
FPS2 overexpression in S. lycopersicum pollen rescued elimination.
- Taxon
- S. lycopersicum
Solanum lycopersicume0 - Treatment
- Overexpression of FPS 2e0
- Gene
- FPS 2e0
- Outcome
- pollen elimination phenotypee0
- Direction
- rescuese0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Overexpression of FPS 2 in S. lycopersicum pollen rescues the pollen elimination phenotype”
Claim · model-written summary
FPS2 pollen selectivity is independent of S-RNase.
- Taxon
- S. lycopersicum
Solanum lycopersicume1 - Gene
- FPS 2e0
- Outcome
- pollen selectivitye0
- Qualifier
- does not involve S‐ RN asee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“FPS 2 ‐based pollen selectivity does not involve S‐ RN ase”
e1“Overexpression of FPS 2 in S. lycopersicum pollen”