Abstract
Aims: Low soil temperature in spring is a major constraint for the cultivation of tropical crops in temperate climates. This study aims at the exploitation of synergistic interactions of micronutrients, consortia of plant growth-promoting microorganisms and N forms as cold-stress protectants. Methods: Maize seedlings were exposed for two weeks to low root zone temperatures at 8-14 °C under controlled conditions on a silty clay-loam soil (pH 6.9) collected from a maize field cultivation site. A pre-selection trial with fungal and bacterial PGPM strains revealed superior cold-protective performance for a microbial consortium of Trichoderma harzianum OMG16 and Bacillus spp. with Zn/Mn supplementation (CombiA+), particularly combined with N-ammonium as a starting point for the characterization of the underlying physiological and molecular mechanisms. Results: In maize nitrate-treated plants, the cold stress treatment increased oxidative leaf damage by 133% and reduced the shoot biomass by 25%, explained by the reduced acquisition of phosphate (P), zinc (Zn) and manganese (Mn). The supplying of N as ammonium improved the Zn and Mn maize nutritional status and increased the ABA shoot concentration by 33%, as well as moderately increased detoxification of reactive oxygen species (ROS). Moreover, use of N as ammonium also increased the root auxin (IAA) concentration (+76%), with increased expression of auxin-responsive genes, involved in IAA synthesis (ZmTSA), transport (ZmPIN1a), and perception (ZmARF12). Additional inoculation with the microbial consortium promoted root colonization with the inoculant strain T. harzianum OMG16 in combination with ammonium fertilization (+140%). An increased ABA/cytokinin ratio and increased concentrations of jasmonic (JA) and salicylic acids (SA) were related to a further increase in enzymatic and non-enzymatic ROS detoxification. Additional supplementation with Zn and Mn further increased shoot IAA, root length and total antioxidants, resulting in the highest shoot biomass production and the lowest leaf damage by oxidative chemical species. Conclusions: Our results suggest the mitigation of cold stress and reduction of stress priming effects on maize plants due to improved ROS detoxification and induction of hormonal stress adaptations rely on the strategic combination of stress-protective nutrients with selected microbial inoculants.
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Claims 15
Claim · model-written summary
Cold stress increased oxidative leaf damage by 133% in nitrate-treated maize.
- Treatment
- the cold stress treatmente1
- Outcome
- oxidative leaf damagee1
- Direction
- increasede1
- Value
- 133%e1
- Qualifier
- nitrate-treated plantse1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“In maize nitrate-treated plants, the cold stress treatment increased oxidative leaf damage by 133% and reduced the shoot biomass by 25%”
Claim · model-written summary
Cold stress reduced shoot biomass by 25% in nitrate-treated maize.
- Treatment
- the cold stress treatmente1
- Outcome
- shoot biomasse1
- Direction
- reducede1
- Value
- 25%e1
- Qualifier
- nitrate-treated plantse1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“In maize nitrate-treated plants, the cold stress treatment increased oxidative leaf damage by 133% and reduced the shoot biomass by 25%”
Claim · model-written summary
The shoot biomass reduction under cold stress is explained by reduced acquisition of phosphate, zinc and manganese.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“In maize nitrate-treated plants, the cold stress treatment increased oxidative leaf damage by 133% and reduced the shoot biomass by 25%”
e2“explained by the reduced acquisition of phosphate (P), zinc (Zn) and manganese (Mn)”
Claim · model-written summary
Supplying N as ammonium improved the zinc and manganese nutritional status of maize.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e3“The supplying of N as ammonium improved the Zn and Mn maize nutritional status and increased the ABA shoot concentration by 33%, as well as moderately increased detoxification of reactive oxygen species (ROS)”
Claim · model-written summary
Supplying N as ammonium increased shoot ABA concentration by 33% in maize.
- Treatment
- supplying of N as ammoniume3
- Compound
- ABAe3
- Outcome
- ABA shoot concentratione3
- Direction
- increasede3
- Value
- 33%e3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e3“The supplying of N as ammonium improved the Zn and Mn maize nutritional status and increased the ABA shoot concentration by 33%, as well as moderately increased detoxification of reactive oxygen species (ROS)”
Claim · model-written summary
Supplying N as ammonium moderately increased detoxification of reactive oxygen species in maize.
- Treatment
- supplying of N as ammoniume3
- Outcome
- detoxification of reactive oxygen species (ROS)e3
- Direction
- increasede3
- Qualifier
- moderatelye3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e3“The supplying of N as ammonium improved the Zn and Mn maize nutritional status and increased the ABA shoot concentration by 33%, as well as moderately increased detoxification of reactive oxygen species (ROS)”
Claim · model-written summary
Use of N as ammonium increased root auxin (IAA) concentration by 76% in maize.
- Treatment
- use of N as ammoniume4
- Outcome
- root auxin (IAA) concentratione4
- Direction
- increasede4
- Value
- +76%e4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e4“use of N as ammonium also increased the root auxin (IAA) concentration (+76%)”
Claim · model-written summary
Use of N as ammonium increased expression of the auxin-responsive genes ZmTSA, ZmPIN1a and ZmARF12 in maize.
- Treatment
- use of N as ammoniume4
- Gene
- ZmTSAe5
- Outcome
- expression of auxin-responsive genese5
- Direction
- increasede5
- Qualifier
- transport (ZmPIN1a), and perception (ZmARF12)e5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e4“use of N as ammonium also increased the root auxin (IAA) concentration (+76%)”
t0“Hormonal Cold Stress Adaptations in Maize”
e5“with increased expression of auxin-responsive genes, involved in IAA synthesis (ZmTSA), transport (ZmPIN1a), and perception (ZmARF12)”
Claim · model-written summary
An increased ABA/cytokinin ratio and higher jasmonic and salicylic acid levels were associated with increased ROS detoxification in maize.
- Compound
- jasmonic (JA) and salicylic acids (SA)e6
- Outcome
- enzymatic and non-enzymatic ROS detoxificatione6
- Direction
- increasee6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e6“An increased ABA/cytokinin ratio and increased concentrations of jasmonic (JA) and salicylic acids (SA) were related to a further increase in enzymatic and non-enzymatic ROS detoxification”
Claim · model-written summary
Additional supplementation with Zn and Mn increased shoot IAA, root length and total antioxidants in maize.
- Treatment
- Additional supplementation with Zn and Mne7
- Outcome
- shoot IAA, root length and total antioxidantse7
- Direction
- increasede7
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e7“Additional supplementation with Zn and Mn further increased shoot IAA, root length and total antioxidants, resulting in the highest shoot biomass production and the lowest leaf damage by oxidative chemical species”
Claim · model-written summary
Additional supplementation with Zn and Mn gave the highest shoot biomass production in maize.
- Treatment
- Additional supplementation with Zn and Mne7
- Outcome
- shoot biomass productione7
- Direction
- higheste7
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e7“Additional supplementation with Zn and Mn further increased shoot IAA, root length and total antioxidants, resulting in the highest shoot biomass production and the lowest leaf damage by oxidative chemical species”
Claim · model-written summary
Additional supplementation with Zn and Mn gave the lowest leaf damage in maize.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e7“Additional supplementation with Zn and Mn further increased shoot IAA, root length and total antioxidants, resulting in the highest shoot biomass production and the lowest leaf damage by oxidative chemical species”
Claim · model-written summary
Inoculation with the microbial consortium promoted root colonization by T. harzianum OMG16 in combination with ammonium fertilization (+140%) in maize.
- Treatment
- inoculation with the microbial consortiume1
- Interactor
- T. harzianum OMG16
Trichoderma harzianume1symbiont - Outcome
- root colonizatione1
- Direction
- promotede1
- Value
- +140%e1
- Qualifier
- in combination with ammonium fertilizatione1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e1“Additional inoculation with the microbial consortium promoted root colonization with the inoculant strain T. harzianum OMG16 in combination with ammonium fertilization (+140%)”
Claim · model-written summary
A pre-selection trial found superior cold-protective performance for a consortium of Trichoderma harzianum OMG16 and Bacillus spp. with Zn/Mn supplementation.
- Treatment
- a microbial consortium of Trichoderma harzianum OMG16 and Bacillus spp. with Zn/Mn supplementation (CombiA+)e1
- Interactor
- Trichoderma harzianum OMG16
Trichoderma harzianume1symbiont - Second interactor
- Bacillus spp.e1symbiont
- Outcome
- cold-protective performancee1
- Direction
- superiore1
- Comparator
- fungal and bacterial PGPM strainse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e1“A pre-selection trial with fungal and bacterial PGPM strains revealed superior cold-protective performance for a microbial consortium of Trichoderma harzianum OMG16 and Bacillus spp. with Zn/Mn supplementation (CombiA+)”
Claim · model-written summary
Mitigation of cold stress in maize relies on a strategic combination of stress-protective nutrients with selected microbial inoculants.
- Outcome
- mitigation of cold stresse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Hormonal Cold Stress Adaptations in Maize”
e1“Our results suggest the mitigation of cold stress and reduction of stress priming effects on maize plants due to improved ROS detoxification and induction of hormonal stress adaptations rely on the strategic combination of stress-protective nutrients with selected microbial inoculants”
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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.