Abstract
For successful double fertilization in flowering plants (angiosperms), pollen tubes deliver 2 nonmotile sperm cells toward female gametes (egg and central cell, respectively). Heatwaves, especially during the reproduction period, threaten male gametophyte (pollen) development, resulting in severe yield losses. Using maize (Zea mays) as a crop and grass model system, we found strong seed set reduction when moderate heat stress was applied for 2 d during the uni- and bicellular stages of pollen development. We show that heat stress accelerates pollen development and impairs pollen germination capabilities when applied at the unicellular stage. Heat stress at the bicellular stage impairs sperm cell development and transport into pollen tubes. To understand the course of the latter defects, we used marker lines and analyzed the transcriptomes of isolated sperm cells. Heat stress affected the expression of genes associated with transcription, RNA processing and translation, DNA replication, and the cell cycle. This included the genes encoding centromeric histone 3 (CENH3) and α-tubulin. Most genes that were misregulated encode proteins involved in the transition from metaphase to anaphase during pollen mitosis II. Heat stress also activated spindle assembly check point and meta- to anaphase transition genes in sperm cells. In summary, misregulation of the identified genes during heat stress at the bicellular stage results in sperm cell development and transport defects ultimately leading to sterility.
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Claims 7
Claim · model-written summary
Two days of moderate heat during uni- and bicellular pollen stages strongly reduced maize seed set.
- Organ / stage
- uni- and bicellular stages of pollen developmente0
- Treatment
- moderate heat stresse0
- Outcome
- seed sete0
- Direction
- reductione0
- Value
- 2 de0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using maize (Zea mays) as a crop and grass model system, we found strong seed set reduction when moderate heat stress was applied for 2 d during the uni- and bicellular stages of pollen development”
Claim · model-written summary
Heat at the unicellular stage accelerated pollen development.
- Organ / stage
- unicellular stagee0
- Treatment
- heat stresse0
- Outcome
- pollen developmente0
- Direction
- acceleratese0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“heat stress accelerates pollen development and impairs pollen germination capabilities when applied at the unicellular stage”
e1“maize (Zea mays)”
Claim · model-written summary
Heat at the unicellular stage impaired pollen germination.
- Organ / stage
- unicellular stagee0
- Treatment
- heat stresse0
- Outcome
- pollen germination capabilitiese0
- Direction
- impairse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“heat stress accelerates pollen development and impairs pollen germination capabilities when applied at the unicellular stage”
e1“maize (Zea mays)”
Claim · model-written summary
Heat at the bicellular stage impaired sperm cell development and transport into pollen tubes.
- Organ / stage
- bicellular stagee0
- Treatment
- Heat stresse0
- Outcome
- sperm cell development and transport into pollen tubese0
- Direction
- impairse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Heat stress at the bicellular stage impairs sperm cell development and transport into pollen tubes”
e1“maize (Zea mays)”
Claim · model-written summary
Heat affected sperm cell expression of genes involved in transcription, RNA processing, translation, DNA replication and the cell cycle.
- Treatment
- Heat stresse0
- Outcome
- expression of genes associated with transcription, RNA processing and translation, DNA replication, and the cell cyclee0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Heat stress affected the expression of genes associated with transcription, RNA processing and translation, DNA replication, and the cell cycle”
e1“maize (Zea mays)”
Claim · model-written summary
Most misregulated sperm cell genes encoded proteins involved in metaphase-to-anaphase transition.
- Outcome
- genes that were misregulatede0
- Qualifier
- transition from metaphase to anaphase during pollen mitosis IIe0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Most genes that were misregulated encode proteins involved in the transition from metaphase to anaphase during pollen mitosis II”
e1“maize (Zea mays)”
Claim · model-written summary
Heat activated spindle assembly checkpoint and metaphase-to-anaphase transition genes in sperm cells.
- Organ / stage
- sperm cellse0
- Treatment
- Heat stresse0
- Outcome
- spindle assembly check point and meta- to anaphase transition genese0
- Direction
- activatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Heat stress also activated spindle assembly check point and meta- to anaphase transition genes in sperm cells”
e1“maize (Zea mays)”
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