Abstract
Both freezing and drought cause cellular dehydration and elicit similar physiological responses in plants, where similar protective compounds are upregulated, including soluble sugars and proteins that help reduce cellular damage. Drought-freeze cross acclimation occurs when drought exposure enhances the freezing tolerance of plants. However, few studies had investigated the reciprocal effects of freezing on drought tolerance, and it was unknown if these interactions could impact plant productivity. Therefore, I examined the effects of freezing on the drought tolerance of individual species and assessed the implications of freeze-drought interactions in an old field community. Poa pratensis was exposed to fall or spring freezing in chambers and a 3-week drought in a greenhouse. Spring-frozen plants had lower reductions in biomass after a summer drought compared to drought-only plants, but this response did not appear to be correlated with the retention of soluble sugars after freezing. Subsequently, I examined freeze-drought responses for a range of herbaceous species: 6 graminoids (Arrhenatherum elatius, Agrostis stolonifera, Bromus inermis, Lolium perenne, Festuca rubra, Poa compressa) and 2 forbs (Securigera varia and Plantago lanceolata). Exposure to a freezing event enhanced the drought tolerance of Agrostis stolonifera, Bromus inermis, Lolium perenne, Poa compressa and Plantago lanceolata, where declines in biomass were lower for freeze-drought plants compared to drought-only plants. These interactions were not correlated to changes in either leaf soluble sugar or protein content before or during the drought. I then collected intact plant-soil mesocosms from an old field and subjected them to spring freezing in chambers and summer drought in the greenhouse. At the functional group level, cross acclimation was observed for legumes; however, for total biomass and nitrogen content in aboveground tissue, cross acclimation was not observed because interactions and additive effects of freezing and drought varied amongst functional groups. Overall, my results highlight the importance of considering plant stress history in experiments, because stressors in different seasons may interact with one another. Nevertheless, cross acclimation responses at the species level may not always scale up to the community or ecosystem scales.
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Claims 13
Claim · model-written summary
Spring-frozen Poa pratensis plants had lower reductions in biomass after a summer drought than drought-only plants.
- Taxon
- Poa pratensis
Poa pratensise1 - Treatment
- Spring-frozen plantse0
- Outcome
- reductions in biomasse0
- Direction
- lowere0
- Comparator
- drought-only plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
.jpg?width=500)
Verbatim
e0“Spring-frozen plants had lower reductions in biomass after a summer drought compared to drought-only plants”
e1“Poa pratensis”
Claim · model-written summary
The lower biomass reduction of spring-frozen Poa pratensis did not appear to be correlated with the retention of soluble sugars after freezing.
- Taxon
- Poa pratensis
Poa pratensise2 - Compound
- soluble sugarse1
- Outcome
- reductions in biomasse0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
.jpg?width=500)
Verbatim
e0“Spring-frozen plants had lower reductions in biomass after a summer drought”
e1“this response did not appear to be correlated with the retention of soluble sugars after freezing”
e2“Poa pratensis”
Claim · model-written summary
A freezing event enhanced the drought tolerance of Agrostis stolonifera: declines in biomass were lower for freeze-drought plants than for drought-only plants.
- Taxon
- Agrostis stolonifera
Agrostis stoloniferae0 - Treatment
- Exposure to a freezing evente0
- Outcome
- declines in biomasse0
- Direction
- lowere0
- Comparator
- drought-only plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exposure to a freezing event enhanced the drought tolerance of Agrostis stolonifera, Bromus inermis, Lolium perenne, Poa compressa and Plantago lanceolata, where declines in biomass were lower for freeze-drought plants compared to drought-only plants”
Claim · model-written summary
A freezing event enhanced the drought tolerance of Bromus inermis: declines in biomass were lower for freeze-drought plants than for drought-only plants.
- Taxon
- Bromus inermis
Bromus inermise0
- Treatment
- Exposure to a freezing evente0
- Outcome
- declines in biomasse0
- Direction
- lowere0
- Comparator
- drought-only plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Exposure to a freezing event enhanced the drought tolerance of Agrostis stolonifera, Bromus inermis, Lolium perenne, Poa compressa and Plantago lanceolata, where declines in biomass were lower for freeze-drought plants compared to drought-only plants”
Claim · model-written summary
A freezing event enhanced the drought tolerance of Lolium perenne: declines in biomass were lower for freeze-drought plants than for drought-only plants.
- Taxon
- Lolium perenne
Lolium perennee0 - Treatment
- Exposure to a freezing evente0
- Outcome
- declines in biomasse0
- Direction
- lowere0
- Comparator
- drought-only plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exposure to a freezing event enhanced the drought tolerance of Agrostis stolonifera, Bromus inermis, Lolium perenne, Poa compressa and Plantago lanceolata, where declines in biomass were lower for freeze-drought plants compared to drought-only plants”
Claim · model-written summary
A freezing event enhanced the drought tolerance of Poa compressa: declines in biomass were lower for freeze-drought plants than for drought-only plants.
- Taxon
- Poa compressa
Poa compressae0 - Treatment
- Exposure to a freezing evente0
- Outcome
- declines in biomasse0
- Direction
- lowere0
- Comparator
- drought-only plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
.jpg?width=500)
Verbatim
e0“Exposure to a freezing event enhanced the drought tolerance of Agrostis stolonifera, Bromus inermis, Lolium perenne, Poa compressa and Plantago lanceolata, where declines in biomass were lower for freeze-drought plants compared to drought-only plants”
Claim · model-written summary
A freezing event enhanced the drought tolerance of Plantago lanceolata: declines in biomass were lower for freeze-drought plants than for drought-only plants.
- Taxon
- Plantago lanceolata
Plantago lanceolatae0 - Treatment
- Exposure to a freezing evente0
- Outcome
- declines in biomasse0
- Direction
- lowere0
- Comparator
- drought-only plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exposure to a freezing event enhanced the drought tolerance of Agrostis stolonifera, Bromus inermis, Lolium perenne, Poa compressa and Plantago lanceolata, where declines in biomass were lower for freeze-drought plants compared to drought-only plants”
Claim · model-written summary
Changes in leaf soluble sugar or protein content before or during the drought did not correlate with the freeze-drought interactions in the herbaceous species.
Not reportedDirection
Verbatim
e0“These interactions were not correlated to changes in either leaf soluble sugar or protein content before or during the drought”
e1“a range of herbaceous species”
Claim · model-written summary
In old-field mesocosms, freeze-drought cross acclimation was observed for legumes at the functional group level.
Verbatim
e0“At the functional group level, cross acclimation was observed for legumes”
Claim · model-written summary
Cross acclimation was not observed for total biomass and aboveground nitrogen content in the old-field mesocosms.
- Outcome
- total biomass and nitrogen content in aboveground tissuee0
- Qualifier
- cross acclimation was not observede0
Not reportedTaxon
Verbatim
e0“for total biomass and nitrogen content in aboveground tissue, cross acclimation was not observed”
Claim · model-written summary
Interactions and additive effects of freezing and drought varied among functional groups in the old-field community.
- Outcome
- interactions and additive effects of freezing and droughte0
- Qualifier
- varied amongst functional groupse0
Not reportedTaxon
Verbatim
e0“interactions and additive effects of freezing and drought varied amongst functional groups”
Claim · model-written summary
Cross acclimation responses at the species level may not always scale up to the community or ecosystem scales.
- Outcome
- cross acclimation responsese0
- Qualifier
- may not always scale up to the community or ecosystem scalese0
Not reportedTaxon
Verbatim
e0“cross acclimation responses at the species level may not always scale up to the community or ecosystem scales”
Claim · model-written summary
The authors recommend considering plant stress history in experiments, because stressors in different seasons may interact.
- Outcome
- considering plant stress history in experimentse0
Not reportedTaxon
Verbatim
e0“importance of considering plant stress history in experiments”
The same questions in other papers
cold · growth · Agrostis stolonifera (creeping bent grass)Positive and negative results
1 in the opposite direction
- negative
cold · growth · Lolium perenne (perennial ryegrass)Positive and negative results
2 papers in the same direction as this paper · 2 in the opposite direction
- no effect
- negative
- positive
1 paper more in this group
See all 5 papers →cold · growth · Bromus inermis (smooth brome grass)Results agree
1 paper in the same direction as this paper
- positive
cold · growth · Plantago lanceolata (narrow-leaved plantain)Results agree
2 papers in the same direction as this paper
- positive
- positive
Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.